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Supplement to Journal of Morphology, Vol. XII, No. 2 



ff(e Stt^eiuntni pttitt 


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ICG/3 77 





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** That which is is far off and exceeding deep : 
who can find it out ? " 

** Because however much a man labour to seek 
it out, yet shall he not find it; yea, moreover, 
though a wise man seek to know it, yet shall 
he not be able." EccUriasU,. 

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Visible Protoplasmic Structure . 

Composition of the Protoplasmic 
Foam 13 

Discontinuous Elements, or In- 
clusions 14 

Continuous, or living, Element 22 

Areal Differentiation based on 

Foam Structure ... 41 Heredity , 

Protoplasmic Activities and Cell- Habit 

Division 67 Instincts 

Striation 89 Conclusion 

Contractility . . .103 

New Structural Formula for Pro- 
toplasm 106 

The Living Substance : As Such 
and as Organism . no 

True Biological Standpoint . 119 

Selection of Environment by the 
Living Substance 

Parasitism . 





The editing of certain studies in protoplasmic phenomena 
having been set aside again and again by circumstances, I am 
impelled to publish a somewhat condensed account of the more 
important facts which bear on problems of the day. 

To do this is to do less justice to the work than one could 
wish, seeing that the larger truths presented were built up as 
mosaics by mere accretion of individual facts ; a method for- 
bidden by the scope of a summary when the number of facts 
is legion. 

Yet it is possible to give a bird's-eye view of the ground 
covered, and, perhaps, to justify those broader statements, 

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which, however they may read, here and there, under such con- 
straint as hypothesis and argument, are but summarized facts. 

The work, at time of its doing, embodied an appeal to the 
normal, living substance, as against '' preserved " and tortured 
states, in which the prevailing standpoint delighted, and from 
which even workers of physiological bias had not freed them- 

As pioneer work it then ran sufficiently counter to prevail- 
ing enthusiasms to be in danger of harsh dealing; to-day it 
may hope for some scattered sympathy. A direct, independent 
appeal to protoplasm, per se, which I have been led to call in 
terms of my results, the living substance, as such; it was unham- 
pered by theory or predilection, except what may be described 
as a belief in life-genii more complex and more potent than 
even surface tension and osmosis. 

The work was carried on partly at the Marine Laboratory at 
Wood's HoU, partly at the University of Pennsylvania, and fin- 
ished in the spring of 1894. It was assisted by constant study, 
during the preceding ten years, of many of the forms then 
used for specialized research. The very full memoranda made 
always at the moment have since been confirmed by frequent 
and iterated observations on the same, or closely allied, mate- 
rial. A few new facts gained in the course of these are omitted 
here, since they are in the nature of confirmation purely, and 
the mass of facts was already too great to be used in toto. 

All controversial references, except a few to Biitschli's epoch- 
making work on protoplasm, to which I owe much, and which 
gave the point of departure for these researches, have been 
omitted, both because the work, save for this, was wholly inde- 
pendent, and because such reference would have increased the 
paper to a size unsuitable even for book form.^ 

And a controversial tone is wholly unnecessary, since the 
facts harmonize rather than clash with all other well-authenti- 
cated facts known to me. 

^ As an understanding of Biitschli's views is advisable for perfect comprehen- 
sion of this paper, it is recommended to those who have not opportunity to read 
his whole work, to read a review of it by Dr. £. A. Andrews ^ich appeared in 
Science, n.s., vol. II, Dec. 27, 1895. ^'^ Biitschli's magnificent work the existing 
theories are reviewed. 

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The new facts are not thought to explain the phenomena, 
but only to unify them, so that the future master who shall find 
for us an explanation may have a less incoherent mass of con- 
tradictory evidences to deal with. 

Though it is felt that acknowledgment to those who have 
from time to time befriended the work by giving opportunities 
to the worker, were better left for a later and more complete 
presentment, which should more fully justify their faith and 
requite their kindness ; it is impossible to omit here recogni- 
tion of help without which the work might never have been 

To Dr. Edmund B. Wilson, who in 1888 admitted me as a 
hearer to his class at Bryn Mawr College, I owe such technical 
training as I possess ; and also then, and later at Wood's HoU, 
stimulus and inspiration to research work, such as springs natu- 
rally from contact with so gifted and catholic a mind, and such 
as grows under an unflagging interest and practical help given 
all who show interest in their work. 

Through the kindness of Dr. C. O. Whitman, I occupied for 
three summers a room in the Investigator's Department of the 
Marine Biological Laboratory at Wood's Holl, Mass., and 
received much encouragement and stimulus, both from him 
personally and from the environment his scientific achieve- 
ments and genial influence create, — a circle of disinterested 
workers who gather about him there, making an atmosphere 
electric with enthusiasm, and holding neither method nor result 
a secret from one another.^ 

Finally, the liberal practical aid of Dr. Horace Jayne, Dean 
of the University of Pennsylvania, providing a private labo- 
ratory in the Biological Department, greatly furthered the 
work and enabled me to bring it,^so far as it goes, to a satis- 
factory conclusion during the winter of 1894. The perfect 
optical conditions enjoyed there, the open north light and free- 
dom from vibrations, gained me some valuable points; and 

^ I desire distmctly to free from all responsibility^ for my choice of a subject, 
or its treatment, the above savants. The privileges and help acknowledged here 
were given to aid research on the embryology of rotifers, the results from which, 
because incomplete, are still unpublished. The present work was carried on at 
the same time and was known to myself only, being then but fragmentary results. 

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happy chance also brought within my reach in the conservatory 
some material specially adapted to certain of the problems. 

Thanks are due also the great courtesy shown by M. Lacaze 
Duthiers, by whose permission I was enabled to enjoy, in the 
summer of 1894, the unique privileges of the Zoological Labo- 
ratory at Roscoflf, France, and there to repeat, under most 
favorable circumstances, the observations on starfish and echi- 
nus eggs; upon which facts, because of their very startling 
nature, I should otherwise have been loath to build from one 
series of observations. 


The optical instruments were, with one exception, of Zeiss ; 
a Zeiss stand, largest size, and accessories ; 4 mm., and 2 mm. 
immersion, objectives; compensating eyepieces, 4, 6, 8, 12. The 
exception spoken of was a picked ^ immersion of Beck's. 
The work done with this was afterward verified with the Zeiss 
tools. The illumination was from an Abb6 condenser, with at 
night blue glass, and a bull's-eye lens, or waxed-paper screen. 

Considering the nature of the work, it should possibly be 
mentioned in this connection that it was further assisted by 
exceptionally far-sighted eyes, having great range and swiftness 
of accommodation. 

The optical conditions were found to be of utmost impor- 
tance. Pure light on a clear day, from a northern exposure, 
gave best results. Failing this, good artificial illumination 
with above conditions was preferred, with precautions against 
undue heat. 

The work done with higher combinations was always com- 
pared, where possible, with results given by a series of lower 
powers ; that done with lowei^powers was verified always under 
higher combinations ; so that optical appearances were thus 
compared with their physical basis of structure. This method 
gave often important results. 

For the more delicate structures and phenomena a sensitive 
adjustment of optical conditions was found to be essential, and 

1 For the reader's convenience the leading results are indicated by numbers in 

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slight variations in these latter, or even in the physiological 
state of the observer, served to hide important facts. 

[i] All observations were made upon the living material, and 
in many instances accompanied by, and compared with, "preser- 
vation methods" of approved sorts. These latter served chiefly 
to show their own inadequacy, acting either as stimuli or as 
relaxers of the substance ; so that, although one might fix beau- 
tiful structures, and even so delicate structures as those figured 
by Biitschli, one had not strictly the original structure, certainly 
not throughout the mass. The impossibility of getting reagents 
to fix simultaneously all parts of even small masses is a great, 
a serious drawback ; even hot osmic fumes failing often to do 
more than create a compound of locally altered states of the 
irritable and active substance. 

I have convinced myself that " preservatives " fix for us little 
of the true structure of the living substance, and can, at best, 
keep for us grosser relations, of a mixed sort in point of time ; 
hiding an infinite complexity of form, and destroying perforce 
those infinitely delicate relations whose fleeting harmonies 
make up life phenomena. I speak of "preservation methods" 
of the past and present, not of the future. 

They have been invaluable in awakening us to the knowledge 
that something we had not known lay hid in the substance. 

The oracle was delivered to us in colored hieroglyphs, to 
which as yet we have no Rosetta stone. Do these speak of 
structural, or of chemical, differences in the living substance ? 

It has been thought they tell of both. Our judgment as to 
the results in individual cases must undergo revision, possibly 
suffer reversion. 

[2] For do such chemical differences as seem to be registered 
pertain to the actual living substance ; or merely to the sub- 
stances mingled with it, and forming the environment it creates 
locally for itself ? 

[3] Again, may not a seeming chemical difference be, per- 
haps, merely a difference of accessibility to these substances, 
grounded on local states, as of viscosity, in a lamellar substance? 

[4] And must we, for such states, appeal to final physical, 
that is, molecular, conditions, or causes ; or can we refer them 

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first to physiological states of the living substance, more within 
the range of our faculties, and capable of being grouped with, 
if not interpreted by, phenomena already known to us? 

These questions are not gropings far afield ; they have not 
till now been asked from the standpoint of the facts gathered 
here, but these force them upon us in just this form and 
through them open up new vistas of research. 

[5] Transcending all the difficulties of preservation, was the 
fact that the protoplasmic structure was found to be in most cases 
an unstable and often evanescent organization of the elements; 
and even where a stable structure existed, such as some of 
those figured for us by Butschli, this was found to be secondary 
in importance to swift and subtle substance-changes which it 

Transmutations, transitions, permutations, metamorphoses: 
— all these, in ** fixed'' material can be, at best, but "fixed " states, 
or appearances, leading to misinterpretation as structural dif- 
ferences, and as such, possible causes even, whereas, in truth, 
they are registered effects of causes which forever defy fixation. 

Special training of the automatic, or registering and will-less, 
attention was devised and found of great use. I believe these 
more purely animal faculties, when properly used, will give 
best results in dealing with such swiftly evanescent phenomena 
as we must learn in protoplasm ; in presence of which a more 
direct exercise of attention by the will is elliptical, confused, 
misled, or baffled utterly. 

Camera drawing of the finest structure and phenomena I 
found impossible. Of the living substance-phenomena, it is 
about as practicable as it would be to trace upon a wall reflec- 
tions thrown there from disturbed water. Some camera draw- 
ings of the filose phenomena in starfish and echinus eggs were 
made, but such can show by direct tracing larger masses only 
of the substance, and are in point of time-relation true but to 
a limited extent. It follows from the very nature of these 
phenomena, as will be seen, that they cannot be traced. For 
while the hand follows one minutest portion, the relations of 
the whole will have undergone change, with important trans- 
positions and transmutations of both structure and substance. 

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In point of minuteness, also, many of the facts which can be 
optically noted defy camera tracing, since the finest steel point 
is still too coarse for the work. 

Visible Protoplasmic Structure. 

[6] The structure of protoplasm throughout the substance 
of all living organisms examined, except where secondarily 
altered, was found to be, as maintained by Butschli, that of a 
visco-fluid foam. 

This is thought to be amply proven by the following optical 
appearances : 

1. By presence of an optical reticulum, such as described by 

2. By presence very commonly, above and below, in proper 
focal relation to this, of a blistered, or what might be termed 
a shagreen-like appearance, which with inadequate powers looks 
slightly granular, or uneven. 

3. Since in all cases, above or below the shagreen surface, 
or the network, proper focus yielded a smooth expanse, variably 
thick, of seemingly homogeneous protoplasm; such expanse 
being large or small according to the amount of the network 
in one optical plane at any moment. 

4. By presence of protoplasmic films or pellicles interven- 
ing between the substance and all contacts involving a 
viscosity different from its own, whether internal or external 
to the mass, whether of alien substances or of its own 

5. By spontaneous formation of like pellicles freely against 
all such differences of contact, whenever and wherever they 

6. By spontaneous vacuolization taking place in even very 
viscous areas ; also in homogeneous-seeming areas. 

7. By similar vacuolization under pressure, or action of re- 
agents ; or as marking relaxed states of the substance ; and in 
states heralding, or following, death. 

8. Finally it is proven by true protoplastic metamorphoses 
and activities, not in fluid areas only, but in those which are, as 

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a whole, very viscid, resistant to pressure, elastic, and even 

The " shagreen " appearance above referred to, when caused 
by Biitschli's structure, is best seen in peripheral areas, espe- 
cially where there is a marked alveolar layer ; but whether this 
latter be present or not, the shagreen effect can be strongly 
marked. It is always to be made out also (if there be no 
optical interference of substances), where alveoli, many or few, 
lie in the same plane, or nearly so. The blistered appearance 
may not come to the surface of pellicles, but lie more or less 
deep within these which vary greatly in thickness; and the 
actual pellicular surface may be covered more or less unstably 
by a far finer shagreen, whose alveoli will measure only from 
one-sixth to one-twelfth /4*. 

Alveolar blistering, whether of the finer froth, or of Biitschli's 
structure, does not always roughen the actual surface, but may 
be much flattened ; and further, may be covered by a secondary 
pellicular film ; or in other cases may project so as to cause an 
optical roughening of the surface when seen from the side. 
I have not, however, in any case yet seen, been able to assure 
myself that the mutual surface of any pellicular shagreen, how- 
ever delicate, as seen from abov^ was not covered by yet 
another pellicle. 

The smooth expanse of protoplasm found everywhere above 
and below networks and alveolar contours proves even more 
conclusively than does the network itself, the presence between 
inclusions of a lamellar substance. In even swiftly flowing and 
unstable areas it can be seen forming structureless, or blank, 
surfaces, which break into fragments the network image, as the 
flux of the substance carries this above or below focus. The 
blank spaces are in their turn broken by rounded contours of 
alveoli rising or falling into focus, and then by reappearance 
of clearly marked network images.' 

1 See Condnuous Substance ; Activities — filose. 

« See p. 9. 

* One may with practice follow for many moments at a time tlie course of small 
groups of alveoli, or even of one or two, in the endosarc of amoebae and other Pro- 
tozoans, and in the early stages of the development of the starfish and echinus 
eggs ; and while doing so, the true nature of these expanses of <' shagreen" texture, 

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Biitschli has not called attention to these film appearances, 
but they are very uniformly present, and to be made out with 
greater or less ease as other conditions may allow. In his 
figures they are probably to be understood in many places 
where mere breaks in the optical network are given. In the 
living substance also, similar breaks in continuity of an optical 
network are seen constantly, without any other image indicating 

[7] Not only is a structure akin to that described and figured 
by Biitschli found, as he asserts, ever)nvhere in living proto- 
plasm; but the facts go even beyond his assertion; for the 
substance of his reticulum, that is the ''lamellar" and pellicular 
substance, is found to be itself alveolated; and to have the 
physical form of a visco-fluid foam. 

This secondary structure is most often visible on pellicular 
surfaces, where it is seen as a delicately blistered, or "sha- 
green" surface, or as delicate striation lines ^; yielding at times, 
with adjusted focus, a network whose meshes are but one- 
sixth to one-twelfth the diameter, as to alveoli, of those of 
Biitschli's structure. A pellicle frequently ofiFers room for sev- 
eral layers of such minute alveoli, yet because of their optical 
smallness, I have not seen in a pellicle, more than one layer.^ 
In the meshes proper, the secondary structure sometimes ap- 
pears as granulation, or vesiculation, or as delicate striation; 
which there, as well as in the pellicles, may be stable, inter- 
mittent, or rhythmical. 

The secondary structure is also seen in protoplasmic films, or 
in extensions from the network of Biitschli, which are, as to 
their whole mass, less than >^ /i. In such cases there is 

or of smooth substance, and of their relation to the network, can be made out 
clearly. The presence of firmly defined fibres or networks in that plane does 
not interfere, from any point of view, with these appearances, since the former 
lie embedded between the films, whether these are thick enough to seem smooth, 
or thin enough to look blbtered. Owing to the greater distinctness, optically, of 
many striae or fibrils, it is necessary, where such are present to use great caution 
in focussing for the film appearances. That is, such as belong to another plane 
of network, for the greater refractiveness common to fibrillar structures may 
optically interfere if they be above the focus. 

^ See Striation. 

> See Activities— filose. 

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often a minute alveolation, which is most unstable as to size 
and arrangement, instead of the much elongated meshes of 
Biitschli's structure, which sometimes take its place. 

Although an optically linear, hexagonal, or stretched reticu- 
lum of Biitschli can be seen in many cases, there are others, 
and not a few, in which this state of things is preceded by the 
alveoli being separated from each other by a considerable amount 
of interalveolar foam; and in place of yielding a true "network" 
they have a very irregular or subspherical contour. They pre- 
sent, in short, just that difference of appearance which is seen 
when the bubbles of an ordinary soap-foam are surrounded by 
a finer froth and are not in direct lamellar contact with other 
bubbles of their own size. 

The local tensions in the fine foam are constantly making 
the outline of the larger bubbles flatten on this side or on that, 
or on several sides at once. The whole seeming of such proto- 
plasmic areas is that of an emulsion rather than that of a true 
foam ; but seeing that the interalveolar stufiF is itself a foam, 
and convertible, and converted into the coarser and more regu- 
lar structure of Butschli, has led me to call the whole compound 
a true foam of varied coarseness. ^ 

Fig. I is a simple, outline diagram of such an area, taken 
from nature, of the protoplasm of a 
portion of a starfish egg, before the 
structure of Biitchli has been more 
perfected. The smaller vesicles show 
the finer foam of the ectosarc-like 
layer, which first forms about the pe- 
riphery of the mass. In this, also, the 
network is broader and more irregular 
than it will be later; and the same 
general characters of the internal area 
are repeated. 

The "granules" in such areas are 
not by any means wholly confined to 
the nodes of the network but are found, as drawn, scattered 
through the finer foam. 

In the endosarc of Vorticellidae, and in their cuticular struc- 

FlG. I. 

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tures during relaxation, such areas are very common. The 
most beautiful and regular structure of this sort I have seen 
was in an immature and unfertilized minnow egg. In the 
protoplasm of Hydra also exist such areas. 

That the appearances, described and figured here, can be 
explained as an optical illusion, grounded in maladjustment of 
focus, it is impossible to allow. The actual thickenings of the 
meshwork, which are common in strial phenomena, are other 
instances and proofs of the existence of a finer foam between 
alveoli of the structure of Biitschli, and with powers inadequate to 
resolve this a greater amount of Continuous Substance than is 
compatible with acceptance of Biitschli's structure as basic can 
be readily seen in innumerable cases. 

The reader is asked to compare this figure with the drawings 
in Biitschli's work from preserved material, and even from life, 
which represent, indeed, veritable and distinct structures of 
Biitschli as they are formed by natural or artificially forced 
organization of the elements. 

The reticulum, then, is not in all, or even in most, cases a 
mere structureless film of homogeneous substance between the 
vesicles of Biitschli's structure; but a compound and highly com- 
plex mass of protoplasmic foam, capable of the same phe- 
nomena as are areas formed of the structure of Biitschli ; which 
latter is often seen to be directly formed, or evolved, from it 
by just such changes.^ 

There are protoplasmic structures and organs, and even or- 
ganisms, whose whole mass is less than that of the interal- 
veolar substance of Biitschli's structure in many cases ; and yet 
within their limits takes place a full series of life activities dif- 
fering in no essential respect from those characterizing large 
masses of living substance. 

Biitschli himself concedes to the structure associated with his 
name considerable range in size, — from j4 to i fi. 

Even in areas where this special structure is marked; and can 
be readily seen to be characteristic; as in Protozoa, and in early 
stages of Metazoan embryos, there are intermixed vesicles of 

^ For full evidence of these assertions, the reader must turn to later portions of 
this paper. 

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all sizes, all gradations existing between a structure of i fi and 
the fine pellicular structure above described, while transmuta- 
tions of one set into the other may be readily seen. 

On the other hand, there are many areas to be found 
where the structure of Butschli is wonderfully perfect and 

[8] In such areas, as in all others whose substance I have 
been able to trace from earlier states, I find that Biitschli's 
structure expresses a secondary and special organization of the 
distribution of the elements of protoplasm, rather than a final, 
or intrinsic, arrangement of these. 

Kindred organization of the elements is seen in the Protozoa 
everywhere, heralding, and indeed making possible, areal spe- 
cialization and areal organization of physiological function ; and 
in the development of Metazoan embryos, the larval and em- 
bryonic physiological areas, or organs, were seen to be laid 
down in the same way.^ 

[9] To sum up : Biitschli's structure, that is, the vesiculation 
of J^ to I ^, is the true structure of protoplasm in so far only 
as it is a characteristic arrangement of the elements in the form 
of a viscous, foam-like, emulsion. 

[10] The secondary finer structures of the meshwork sub- 
stance, taken in connection with certain activities of this, to be 
described further on, seem to place beyond dispute, that the 
structure of J^ to i ft, is not, indeed, the final structure of the 
living substance, but that it is part only of an infinitely graded 
series of vesiculations of the protoplasmic foam ; whose minute- 
ness it would be rash to limit even to the finest pellicular net- 
work visible ; and whose coarseness it would be still more 
arbitrary to limit by one ft. 

These facts were established during research upon the follow- 
ing organisms : Amoeba proteus. Amoeba radiosa ; Actinophrys 
sol ; Actinosphaerium Eichomii ; Raphidiophrys, of several un- 
identified species ; Monads of numerous sorts ; Choano-Flagel- 
lates of several species ; Vorticellidae in several families, notably 
Epistylis ; Myxomycete found in salt water, Myxomycete found 
in fresh water ; Stentor ; Acinetans ; Coleps hirtus ; and many 

1 See Areal Differentiation, and Striation. 

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Other forms of the Protozoa which crossed the path of my work 
from time to time, and were used passingly as corroborative 
evidence ; Hydra viridis and Hydra f usca ; Turbellarians ; An- 
guillulidae ; Rotif eras, adult and embryo ; Starfish and Echinus 
embryos and larvae ; frogs' eggs ; fish eggs ; tadpole's tissues ; 
crabs' tissues and the Leucocytes of crabs' blood ; also many 
fresh-water Crustacea; fresh sciatic muscle of the dog. In 
plants, Spirogyra, Chara, Nitella. 

Composition of the Protoplasmic Foam, 

The protoplasmic froth is infinitely complex in character of its 
constituents, and in arrangement and grouping of these, which, 
in Metazoa as well as in Protozoa, are subject to more or less 
gradual change of size, and of actual, as well as relative, posi- 

[11] A substance formulated as a visco-fluid foam must have 
its elements, however multiple their character, divided at any 
given moment into two broad physical groups. This is the 
state of things we find characterizing protoplasm. 

There is first the element, be its nature simple or complex, 
which forms at any given moment the lamellar, or enclosing 
substance, of the foam vesicles, throughout the whole or any 
part of the series of structural subdivisions. 

This is, physically, the Continuous Substance. 

Secondly, we have those elements seen as the substances in- 
cluded in the vacuoles, or cavities, of the foam : these are isolated 
to some extent from each other by lamellar films ; and in each 
structural series by interalveolar material, which in different 
masses or areas, or in the same areas from moment to moment, 
varies in thickness, in constitution, and structure, as well as in 
quality, or viscidity. 

The substances so isolated are, therefore, at any given moment 
the Inclusions, or the Discontinuous Substances. 

Where the continuous, or interalveolar substance, is itself a 
compound emulsive froth, repeating the characters of the whole 
mass, it is true the two groups of elements, from the standpoint 
of Biitschli's structure, may seem to be in a sense almost inex- 

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tricably mingled ; yet for the physicist they remain clearly dis- 
tinguishable at any given moment. 

That this is true for the physiologist also wUl be presently 


In character these are most heterogeneous. They may be 
subdivided into Simple and Compound inclusions. They are 
fluids, of various degrees of viscidity, with or without suspended 
substances^; or solids; they may be concretions, or secretions, 
or excretions ; native or adventitious. 

Even very small areas are found to contain many chemically 
difiFerent inclusions. 

Under Simple Inclusions are grouped the alveolar contents 
of Biitschli's structure, or of the finer froth, or of coarser alveo- 
lation; wherever no interpenetration of the continuous element, 
subdividing the mass into a finer froth can directly or indirectly 
be detected. Simple inclusions may be of one substance, or of 
a number of mechanically mingled substances. They are some- 
times transformed into members of the second group by activi- 
ties of the lamellar substance.^ 

Under the head of Compound Inclusions are placed those 
areas, irrespective of both actual and relative size^ where are 
grouped together alveoli, containing, as far as can be optically 
or chemically demonstrated, the same, or very similar, discon- 
tinuous substances. 

The lamellar substance of these inclusions unites to form one 
common pellicle which, with or without modification of various 
sorts, separates them as a mass from adjacent protoplasm. 
Such groups may be built up on a basis of the finer froth; and 
then form one or more inclusions of Biitschli's structure ; or 
they may be built up of the latter only, or of both ; but in all 
cases the mass is an aggregate of single vesicles, and therefore 
interpenetrated at some time, if not always, by the continuous 
substance, which when visible forms an optical network. 

^ The Protozoa furnish endless instances of mingled indttsiont. 
> See p. 19. 

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Some compound inclusions have become very familiar to us 
under the names of starch, yolk, pigment, the " gjranules " of 
the network ; and chromatin, as fibres, granules, masses, or 

Yet even of these long-known substances much remains to 
be learned. Many questions that now leap into almost impera- 
tive urgency have, up to this time, scarcely shaped themselves. 

Which of these substances are stimuli ; which are food for 
the final protoplasm ; which are excretions, which secretions ; 
which are means to an end ; which attained ends ? In other 
words, which are final, which intermediate, agents in producing 
the physiological results we can trace ? 

Such questions are puzzling enough when applied to the ele- 
ments of the structure of Biitschli. More puzzling still must 
it be to answer which of our results, in any given area, are due 
to the inclusions of this alone ; which to the interalveolar sub- 
stance, with its inclusions. And finally, may not some of the 
results had in the inclusions of the structure of Biitschli, be 
due to secretions or excretions of the interalveolar substance 
under irritation } 

The substance called yolk seems to me one of the most inter- 
esting problems. When one considers that in cases such as 
are met with in certain rotifers, the whole of a huge ovary, con- 
taining material for a score of eggs, is converted, under certain 
conditions, into a single egg ; and that nuclei, as well as the 
already prepared yolk, go to form the " yolk " for this monster 
egg: the question naturally arises; which part of the whole mass 
of intermingled material goes to make up the organism which 
issues forth in the following spring } 

We know that yolk disappears largely, if not wholly, during 
process of development of the immature creature. But let me 
emphasize here what all the facts collected are ceaselessly 
emphasizing: that optical disappearance, under such conditions 
as exist in the protoplasmic froth, may mean several widely 
distinct sets of phenomena. 

Given the finer structure of the interalveolar substance, 
many obscure phenomena associated with the life history of all 
these substances become more intelligible. 

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The structure and growth of starch grains, and possibly 
the behavior of their different layers in swelling ; the froth-like 
structure which no treatment seems to take from them ; the 
redistribution at times of their substance through the proto- 
plasm; the formation and dissolution of yolk, which in some 
cases at least, is a physical process before it seemingly becomes 
a chemical process ; the numerous redistributions seen to take 
place in the nuclear element we call chromatin : — all these 
become intelligible if interpreted as phenomena of the finer 
froth; differing in point of minuteness only from characteristic 
phenomena seen in Biitschli's structure. 

Such an interpretation is not only plausible but is forced 
upon us in many cases by the optical phenomena attending the 
physical fact. It is indeed hardly to be escaped, for down to 
the limits of vision there is found in all parts of living masses 
a certain absolute unity of structure and of phenomena.^ 

[i 2] Similar evidences that the network ^^ granules " are of the 
nature of compound inclusions, are to be had in their disappear- 
ance during structural reductions, or subdivisions of Biitschli's 
structure, which produce an appearance of homogeneous proto- 
plasm ; and in their reappearance upon return of the structure 
to the first coarser organization which produced a distinct 

[13] The universal presence of the "granules," or of the ele- 
ment which is so named, as seen in Biitschli's network, suggests 
that we have here an element, which is in some way essential to 
some if not all of the characteristic activities of the substance ; 
that in the form in which it is best known it stands not merely 
for a passive chemical inclusion, but for a physiological area or 
substance organ.* 

Certain strong resemblances of the granules to the chromatin 
of nuclear threads may be not wholly superficial, but hold the clue 
to a right interpretation of these most interesting substances.^ 

How long the lamellar substance of compound inclusions 
persists in a living state, and what, during that time, may be 

^ See Areal Differentiation ; also Activities — filose. 

* Idem ; also Habit. 

* For movements of the granules, see Activities — filose. 

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its relations with the surrounding protoplasm ; whether, unlike 
the major part of the network stuff, it is stable in position ; 
or like this, in constant flux of intercourse with surrounding 
areas ; whether the inclusions are in constant use by the sub- 
stance ; or whether they are simply storehouses of chemical in- 
fluence, withheld for special times or crises in the life-rhythms 
of the mass, or portions of it : — all these questions now present 
themselves as to the substances distinguished in Biitschli's 
structure ; and we need the answer. 

Inclusions may also be subdivided into microscopic and macro- 
scopic. Microscopic^ are those which lie within the lamellar 
alveoli of Biitschli's structure, or of the finer froth ; macroscopic^ 
those which are built up of these, forming, in any living mass, 
areas, or structures, relatively large. 

[14] When dealing with the living substance strictly as 
such, and with organisms as masses of protoplasmic foam; 
all endo-skeletons, all vascular systems, all body cavities, 
have for the substance, or the mass, the same value in a 
physical sense as the alveolar inclusions have for the lamel- 
lar substance. 

Even exo-skeletons arise in many cases as protoplasmic in- 
clusions, and later, by withdrawal, or atrophy, of the living sub- 
stance, become more strictly external. 

It is interesting that this statement, in at least one instance, 
can be applied to the membrane of plants also. The cellulose 
wall of very young Spirogyra filaments was seen to be covered 
by a most delicate film of living substance, of which the alveoli 
were hardly one-sixth the size of alveoli of Biitschli's structure. 
They were elongated and arranged in parallel rows. Most 
curious of all was it that the rows of alveoli followed with 
perfect sympathy the direction of the jntemal spirally twisted 
chlorophyll bands, so that the surface under a lower power 
appeared most delicately striated longitudinally; as if there 
were in the nature of the whole plant substance the same 
spiral tendencies. The inclusions appeared to be homogene- 
ously fluid. In older specimens this outer film disappears; 
either obliterated by the increasing deposits of cellulose, or 
having itself migrated to the interior of the cell. The 

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cellulose wall of Spirogyra, then, exists at first as a sub- 
stance inclusion.^ 

Throughout the series of structural subdivisions of proto- 
plasm the inclusions are for the most part unstable. They are 
seen to increase either slowly or rapidly by purely physical 
changes ; and they suffer change also by what must still be 
called physiological activities of the interalveolar protoplasm. 
They grow by absorption of material from, or through, the sur- 
rounding substance ; they wane by diffusion in the same mode ; 
they are at times suddenly augmented by relaxed states of the 
nearest interalveolar stuff, or of the whole area, or mass, and may 
then attain such size as to rank with " vacuolations "; or, from 
a size too small to be directly seen as alveoli, may be brought 
into the range of the structure of Biitschli. 

Inclusions may be also more or less rapidly increased in size 
by withdrawal of the interalveolar substance, which involves, 
or leads to bursting of, the lamellae ; throwing together the 
contents of adjoining alveoli. Besides a purely physical cause 
of this sort, such as is familiar in soap-foams, there are cases 
in which the interalveolar, if not the lamellar, substance craivls 
or flows away from between alveoli. This phenomenon has 
been seen in many Protozoa; typical instances are readily 
watched in Heliozoa, as Actinosphaerium. In the formation by 
these of tubular rays, and by Acinetans of hollow tubes or suc- 
torial tentacles, there can be little doubt as to the way in which 
the transfer of protoplasm takes place.^ 

In Acinetans there is formed a rod-like extension of the 
substance, which by successive redistributions of the elements 
becomes at last a mere row of alveoli of about equal size. The 
network of the whole has the effect of a "ladder,*' whose 
sides are increasingly thickened by immigration of protoplasm 
from the "rounds." These latter are finally obliterated as a 
result of this, and the whole interior is then one long closed 

^ It is not impossible that to contractions of such a film of organized proto- 
plasm may be due the movements of many plant filaments, as Oscillatoria, etc. 
See Striation. 

3 Each one of these phenomena, it must be remembered, can be fully grasped 
only by aid of all the facts given in this paper, which are supported by hundreds 
not given. For above, see also Activities of the Continuous Substance. 

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tube full of fluid which has a pinkish tint. By similar activi- 
ties the end of the tube becomes very much thinned, and at last 
bursts open, much as a bubble would, in the centre. The proto- 
plasm then moulds itself about the rim and the tentacle is 
ready for use. The whole process occupies a few moments. 

This is one of numberless instances, which go to show that 
it is the interalveolar rather than the inclusion substance of 
protoplasmic foams, which is the living substance. 

Fluid alveoli of Biitschli's structure are subdivided in a num- 
ber of ways. They have been seen to be constricted^ orpinched^ 
in twOf precisely as large vacuoles are at times. This occurs 
in the endosarc of amoebae ; in the formation of ectosarcal parts 
and organs, and in other reorganizations of the elements. 
They are broken up also by what appears to be a relaxation of 
their lamellar substance, at the same time with the surround- 
ing substance; for this latter at such times seems to lose in vis- 
cidity and to become more or less mechanically vacuolated. 

They are at times penetrated by filose processes spun into 
their interior from the interalveolar substance. Whether the 
actual lamellar substance takes any active part in these proc- 
esses could not of course be optically determined. In the 
endosarc of Amoebae, where the phenomena were first ob- 
served, the simple filose processes were seen to branch. In 
the Vorticellid, Epistylis — in the somewhat larger alveoli of 
whose cuticle the same thing was later seen — the simple filose 
extensions were swiftly ramified, until a distinct network was 
formed. From this, in some cases, a finer froth appeared to 
be formed later, in what manner could only be conjectured. 
The filose processes may spin quite across the alveolus and 
become fused with the opposite wall, or return to the main 
thread, or the wall whence they came. The process is best 
described by likening it to characteristic activities of the Pro- 
toplast Gromia, 

It is not impossible that there may exist already in the fluid 
inclusion a fine network, invisible by reason of the tenuity of 
the lamellar substance, and that the filose processes merely fol- 
low the course of the lamellae, pushing between the yielding 
vesicles very readily, by reason of a rather greater amount of 

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viscidity. That the filose formations were free is not, on the 
other handy at all less likely ^ as will appear under Protoplasmic 
Activities. The phenomena were seen to take place while the 
general cuticle, as well as the whole mass of the animal, was 
quite strongly contracted. 

[15] This proved that under the mask of a stable and even a 
contracted area of Butschli's structure, local relaxations may 
exist, rendering possible not only transfer of substance but 
modification of the finer structure of the living substance. It 
also went to show that of the two groups of elements it is not 
the alveolar contents which play the leading rdle.^ 

Variations of size of alveoli in correlation with physiologfical 
function will be discussed further on.^ 

[16] There are found in the living substance innumerable 
contradictions of the conditions ruling the artificial foams of 
Biitschli. Many of these are to be cited. It may be stated 
here that as to size and distribution of inclusions, as in most 
other particulars, while there are always cases to be met with 
which coincide with the inorganic foams, there are more which 
cannot be made to agree. 

An adequate explanation of living phenomena must meet all 
the facts. 

Such intermingling of various matters as exists in protoplasm 
must render chemical analysis of this, as living substance y forever 
difficult, for besides the fact that the two sets of elements are 
so finely subdivided and so intermixed in very small portions, — 
portions so small indeed as to pass at times beyond microscopical 
vision,^ — how shall it be determined which part of the result 
belongs to Which group } In our eflForts to learn something 
of the actual living substance, how shall we know which part 
of a given reaction, or which among all the detected qualities 
or matter, belong indeed to the living element ; which to the 
intermingled and non-living matter } How may we know, which 
part of local or temporary variations are due to local variations in 
these latter; which to physiological difiFerence in the functioning 

1 See Activities — filose. Also Striation. 
' See Areal Differentiation — Ectosarc. 
* See Living Substance as Such. 

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substance ? And finally, which part of the constant results is 
constant as living substance ; which but as its environment ? 

A still more radical question has not yet been disposed of : 
how are we to know that there is in the whole compound 
anything more than interaction, under physical controls, bom 
of their physical arrangement; of non-living, of chemical 
materials ? 

All of these questions, not excluding the last, have received 
for me illumination from study of the inclusions of living proto- 
plasm ; and yet more light from study of the continuous sub- 
stance. Though not answered, they receive limitation which 
may later help us to an answer. 

Nothing is more common in protoplasm than an areal organi- 
zation, more or less unstable, of the vital phenomena, so that 
such areas are traversed by the same special phenomena, and 
the whole may be called physiologically homogeneous through- 
out. Yet in such area of organized physiological function, the 
inclusions of Biitschli may or may not show homogeneity in 
kind or size, but may be thoroughly heterogeneous.^ 

While this is possibly explicable by supposing homogene- 
ousness in character of the finer froth inclusions, this is but to 
emphasize the answer already indicated ; and further, to show 
that not only must we not look to the inclusions of Biitschli's 
structure for the basis of such homogeneity of function ; but 
that we shall find it in the interalveolar stufiF. 

A sensitively organized activity, and reaction to stimuli, 
throughout an area whose inclusions are mingled solids and 
fluids, and of all sizes, may hardly be interpreted as physical 
interaction between a homogeneous lamellar substance and such 
chemical and physical heterogeneity. Neither diffusion nor 
surface tension can readily be brought into play here. On the 
other hand, the continuous substance in such areas, as in all 
others where organized physiological activities are found, has 
definite and most homogeneous characters. 

[17] In the section on contractility it will be shown that in 
this most important activity, so far as all optical evidence goes, 
the alveolar contents form the passive group of elements, 

1 See Striation — fibrillar. 

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responsive by change of shape or relative position to compulsion 
of interalveolar activities ; while the network substance, down 
to its most minute subdivision, shows a true, characteristic 
rhythm of shape and position.^ 

[i8] It is the mode of distribution and optical characters of 
the continuous substance, as well as certain constant changes 
that take place in these, which further make it certain that 
in the structure of Biitschli, and beyond this to the limit of 
vision, and again beyond this, it is the continuous substance 
which manifests the characteristic activities of the substance, 

[19] To sum up: there was found no optical evidence in any 
material examined, to show that the part played by the discon- 
tinuous substances is anything more than a passive chemical 
and physical r61e ; while there was endless and ever unfolding 
evidence that it is the continuous substance, at any given 
moment, which sustains the rdle of the living protoplasm ; and 
as far down in the scale as the human eye with its most power- 
ful aids can trace it, it is this same continuous substance which 
we must be content yet awhile to call the living, the physiologi- 
cally active, stuff. 


In dealing with the coihinuous element it becomes neces- 
sary, in view of its compound nature, to subdivide verbally 
what has hitherto been spoken of as Biitschli's lamellar sub- 
stance, or Biitschli's network. It is necessary to distinguish in 
various cases the whole continuous substance of the physicist, 
which includes in a given structure both alveolar lamellae and 
the interalveolar foam ; from the latter as a thing by itself. 
This is the more forced upon us since we must now learn to 
think of optical networks and lines as by no means identical 
with those actual fibrils and networks whose existence is shortly 
to receive proof. 

The word " network," as commonly used, brings with it an 
element of confusion, in that it does not clearly distinguish 

^ See Striation and Contractility. 

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between actual lines and included spaces. Further *« network ** 
ill expresses a structure existing in three dimensions of space. 

I shall therefore restrict the use of this word to the optical 
network effect, whether this at the moment expresses a physi- 
cal fact or an optical figure; and also, when not otherwise 
specified, to mean the intersecting lines of this, excluding 

By continuous substance will be meant the whole substance 
separating inclusions — that is, it will be used in the strict 
physical sense. 

Interalveolar stuffy or material or foanty or substance^ or pro- 
toplasm, will be used for the continuous substance, exclusive of 
the actual alveolar lamella : and interalveolar structure y as de- 
scriptive of the distribution of the two sets of elements in this 

By this phraseology, which will presently be seen to be very 
necessarily precise, many mental difficulties and much vague- 
ness are evaded in dealing with protoplasmic foams. 


Our conception of the lamellar substance of Biitschli's struc- 
ture being enlarged by knowledge of the finer froth structure 
within it, we must now image to ourselves the true continuous 
substance as forming the lamellar and also interalveolar stuff 
of the finer foam ; then, as being continuous by way of these 
with the lamellae and interalveolar stuff of Biitschli's structure ; 
then, by way of these, continuous with all pellicles internal 
to the mass ; and finally, in the same manner, continuous with 
the external, or mass, pellicle and its products. 

This state of things can be seen in many transparent areas, 
and in some entire masses of protoplasm. Those optical appear- 
ances, upon which objection to the fluid foam structure of proto- 
plasm has been based, such as fibrils and networks of indubi- 
table firmness, are here for the moment set aside, as, though 
their existence will later be finally proven, they will also be 
shown to be coexistent with, and in no sense inhibitive of, a 
true foam state. 

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[20] In one respect the kinship between protoplasm and arti- 
ficial foams, as established by Biitschli's masterly researches, 
holds good and is not to be shaken. As to the formation of 
pellicles, the physical nature of the substance seems to rule, and 
never, so far as I have seen, is it set aside by those controls which 
elsewhere appear to dominate from time to time that physical 
mastership of the elements Biitschli has demonstrated. 

This fact, as pointed out above, is one of the strongest evi- 
dences of the true nature of the substance. 

From mass pellicles in all their varieties,^ through the series 
of inclusion pellicles down to alveolar lamellae and pellicles of 
the finer foam inclusions; the living substance maintains its 
habit of surrounding all physical differences of contact with a 
continuous film of its own material. 

But this basic unity of structural form covers an infinity of 
structural differences, of which many organisms; not less those 
which for many years biologists have termed " simple," than 
members of the highest groups ; will furnish abundant proof. 

Only the most obvious of these have so far been noted : 
vast fields for research are now seen to open out in respect to 
those which are not only less conspicuous but also in many 
instances more evanescent organizations of the elements. 

In an amoeba, for instance, the peripheral pellicle which 
stands for cell wall, the ectosarc possibly, the contractile vac- 
uole, the food sacs, the nucleus ; — these have been taken to 
be the limit of organization of the substance in this lump of 
" primitive protoplasm." 

So far as these well-known areas of differentiation of the 
living substance go, they were seen, with respect to the foam 
structure, to be optically separated from each other by charac- 
ter of their inclusions and of the continuous element ; and by 
the mode of distribution of these two in relation to each other, 
not only as permanent differences but as differences from 
moment to moment. 

Besides the broad areal differences noted, there exist local 
differentiations more or less unstable, to the point of such 
evanescence often as makes it difficult to catch a glimpse of 

1 See also Ectosarc. 

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— —1 


their fleeting existence. But none the less are they ureal differ- 
entiations of structure ; and none the less are they associated with 
physiological function of the substance. 

In Metazoan eggs, as of starfish, echinus, rotifer, flsh, or 
frog, in Metazoan larvae even, the same holds equally true. 

It may be said for living protoplasm, that of making pellicles 
there is no end. Yet the pellicles when made differ one from 
the other as markedly as, in many cases, do nuclei, or cell 
membranes, or skeletons, or any other coarser, complex structure 
of the substance. 

The optical network of Biitschli's structure is found in all 
protoplasmic masses I have seen, to be locally emphasized so as 
to form optically greater separation between areas of the sub- 
stance by marked partition lines, plates, or membrane-like thick- 
enings of the lamellar film. 

Such modifications are to be grouped, both in structure and 
origin, with the pellicles surrounding external masses and 
forming contact surfaces. 

Like these, they are actual thickenings of interalveolar stuff 
along lines of the physical lamellae, and mean that here the 
mass of interalveolar foam has been augmented, or has under- 
gone physical modification of some sort. 

We must now begin to distinguish verbally between the 
necessary physical, or purely physical, pellicle and such thicken- 
ings as these. For the former I shall say physical pellicle, or 
lamellar substance; for the latter merely pellicle, or pellicular 
plate, thickening, or membrane. 

[21] Wherever a finer foam structure can, directly or indi- 
rectly, be traced in a pellicle, it follows, from the nature of the 
case, that the outer surface of that pellicle is in its turn covered 
by a pellicle. Hence, wherever pellicles are augmented by 
interalveolar material they are double ; and actual contact with 
environment, whether this be internal, or external, to the mass, 
is made by the pellicle of the finer froth. 

Fine instances of this are seen in the pellicular formations of 

Because of phenomena found associated with flux of pellicu- 
lar substance, it becomes impossible to set a limit to such 

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multiplication of pellicles, even to the end of microscopic 


[22] Pellicles differ one from the other, or at different 
moments of time, from their own former states; in point of 
thickness, of viscidity, of finer structure, and of stability in the 
component protoplasm, as well as in activities of this. The 
same is true of the interalveolar substance throughout its 

The varying viscosity of pellicles is shown by optical increase 
of density, which also brings a characteristic change of hue ; 
and by change of refractive value; with increased resistance to 
pressure and increased elasticity. At such times, owing to 
these optical qualities, structures so minute, or expanses of 
substance so tenuous, as to be in more fluid states wholly 
invisible, become readily detected.* 

It has been urged that to be capable of contraction, proto- 
plasm must needs be a solid substance ; it has also been 
affirmed that if a fluid it cannot be elastic as contractile sub- 
stances are. 

That both of these conditions are fulfilled by the substance 
and that it yet does not for these renounce its typical fluidity 
can be proven by hundreds of instances. A single instance 
will suffice to make this clear. 

The pellicle of the contractile vacuole of amoeba, which is 
at times so fluid as to mix freely with the surrounding proto- 
plasm, becomes at other moments, notably while in that very 
viscid refractive state which rhythmically precedes the collapse 
of the organ ; so very firm and so truly elastic that the vacuole 
suffers constant change of shape under pressure of the endo- 
sarc, of the included substances also, and the peri-neuclear area, 
if present. At such times it looks most like a transparent 
rubber bag filled with water. 

Conversely, the instant return of a most elastic and highly 
organized area to a truly fluid state is beautifully shown, when, 
by breaking or tearing tension, the stalk of a retractile vorti- 
cellid is torn apart. Then the contractile muscular fibre, which 

* See Activities — filose ; also Substance as Such, description of Choano- 

s See Striation. 

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is composed of a number of different areas, suddenly rounds 
itself off and shows a smooth surface at the broken ends, while 
at the same time the different areas fuse smoothly together 
at this region and can no longer be distinguished apart. The 
same experiment can be tried in higher muscular areas, as of 

In relaxed states, areas of the structure of Biitschli which 
were before very viscid, become so fluid in seeming with respect 
to that structure, that the protoplasm presents the appearance 
of mixing freely with water. So with the finer structure; 
states of such viscidity as caused elasticity of the continuous 
or interalveolar element, may be followed by relaxed states in 
which the protoplasm proper of Butschli's structure seems 
miscible with water. 

Such appearances as described above may characterize the 
whole of a pellicular area or be distributed in lines or networks 
within it, at varying distances from the surface. In these latter 
forms, they furnish, at times, the only direct evidence of alveolar 
structure; and they then prove also that there is organization 
of the elements for physiological function.^ 

Upon formation of physical pellicles where the area of con- 
tact, internal or external, involves a number of alveoli, there 
seems to be a tendency in protoplasm to thicken the film so 
formed by access of interalveolar material. 

In a similar manner, that is, by interalveolar material and out- 
flow of what has been termed hyaloplasm from the endosarc, 
ectosarc is formed. Almost all pellicles, certainly all those of 
Biitschli's structure, are in strict sense an ectosarcal formation, 
no matter what position they hold in the mass. 

The viscosity of the continuous substance varies locally from 
moment to moment, the changes being often miraculously 
swift. From a very fluid state it will become rapidly so viscous 
as to resist pressure after the manner of a stiff elastic bristle ; 
and this may take place without change in size of the visible 
foam structure. 

In such states it is as a rule markedly contractile; and com- 
monly the assiTmption of organized contractile activities by an 

1 See Striation. 

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area can be foretold by just these characteristic differences, 
which are identical with those seen in most ectosarcal areas 
and products. 

It is by local differences in viscosity that differences of shape 
in masses, which had their origin in protoplasmic activities, 
are maintained, until they may or may not be released to pass 
into physical control of secreted, or adventitious, non-living 
materials. The same is true of those internal cavities, or 
vesicles, whose complex, non-spherical forms have been used 
as an argument against Biitschli's foam theory. 

Variations in viscidity of the substance as masses, or areas, 
are optically manifest as differences in the same manner as 
shown for pellicular substance. 

Increased viscidity does not always mean structural subdi- 
vision of Biitschli's structure, although this of itself is physi- 
cally a common cause of increased viscidity (ceteris paribus) 
wherever it occurs. 

Nor does it necessarily mean in living foams, as it usually 
does in physical foams, an increased tenuity of the lamellar 
membranes of Biitschli's or other visible structure. 

In living foams, the structure of a given area of great 
viscidity may be in size not at all different from surrounding 
areas ; yet the viscidity shown under pressure, or osmotic con- 
ditions, may be far above that of these latter. 

The continuous substance may be no thinner, may even be 
markedly thicker, than that of the adjacent protoplasm, and yet 
the areal viscidity vastly greater. 

Hence comes it, that there is often found an areal viscidity 
which may have almost any structural character in point of size 
of alveoli, or distribution of the continuous element of a given 

All such states and conditions may be seen freely inter- 
changed and playing freely into each other in the protoplast, 
Amoeba radiosa; coarse and fluid structure changing in a 
moment's time into minute and highly refractive areas; 
these returning to coarse and highly viscid states; these 
again to finely subdivided states, which may be either fluid 
or viscid. 

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At one moment the substance becomes a film so tenuous as 
almost and then quite to elude the most careful search, yet 
having a true structure of Biitschli ; then gathers itself up 
into blunt pseudopodia of great thickness, and of double, or 
triple, or manifold, alveolar differentiation ; then spins itself 
out into long delicate whip-lashes, which are at first coarsely 
structured near their bases, but pass into invisibly subdivided 
states, and become so viscid as to bend under mechanical 
pressure of the cover-glass, like a stiff bristle as they extend 
straight out in the water. The tip of these rays may elongate 
still further till a fine thread forms. Before, or after, this last 
change they are optically so refractive that they glitter, and 
have a greenish hue like glass rods. 

In pseudopodia so modified, are seen contractile activities ; 
for they bend about like tactile organs, as, indeed, they prove 
themselves in various ways to be for the time. At other 
moments they bend their length into contorted spirals and 
ram*s-hom-like shapes, and then again, lengthening, will lash 
the water about in a frenzied way exactly like overgrown cilia 
or flagella. 

A momentary touch upon the cover-glass will in one moment 
convert all of this display into inactivity, leaving but a shape- 
less lump of slightly amoeboid protoplasm, which usually sinks 
more or less in the water. That area in the mass from 
whence arose the contractile processes is different for a few 
moments, has a more uneven and disturbed appearance than 
the rest of the lump. And when the creature resumes its 
activities, it is from this same area that there usually arise 
again new organs having the same peculiar characters and 
showing the same metamorphoses and activities ; while those 
portions whose processes were more ordinary pseudopodia again 
produce these ; showing that the activities, though intermitted, 
were only suspended, and that their suspension was a phenom- 
enon physiological, and not of physical control alone. 

It must be remembered that the slight jar to the cover- 
glass was only one of countless vibration disturbances to which 
the animal is subjected ; of a different character from those 
occasioned by passing organisms, but still only a jar, which 

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seems from a physical standpoint inadequate to cause such 
transformation, unless, indeed, it were sufficiently disintegrating 
to prevent renewed manifestations. 

In the vibratile state of the processes, delicate transverse 
striations could fleetingly be seen in their lower portions. 
Delicate, cilia-like, appendages were at times produced from 
very blunt ends of other lobose pseudopodia in this form, and 
were also vibratile, and acted as tactile organs. 

To the student of the substance, this form is recommended 
as uniquely fruitful. 

[23] In all the above facts, areal viscidity of the living sub- 
stance was seen to be something independent of the physical 
distribution of the elements in any given, visible, structure, 
especially if that structure were Biitschli's. 

[24] It is worthy of special note that in some viscid states, 
which show increased elasticity, or even mere increase of 
refraction, pellicles resist longer the passage through them of 
hardening, killing, or even staining, reagents. Their kinship 
with ectosarcal formations is by this more fully established. 
The same property characterized the interalveolar stuff wher- 
ever found in a similar state. Unstable interalveolar formations 
having this quality persisted also in preserved material as distinct 
substance structures, and then had, every appearance of those 
substance structures which are more stable in life. 

These facts offer a valuable hint as to a possible cause of 
such structures as "achromatic protoplasm," or " archoplasm," 
in eggs ; and as to differing results given by preservation 
methods in nuclear and cytoplasmic phenomena at different 
times in the rhythms of cell division phenomena. 

I find that where very viscous states of the interalveolar 
material caused viscid states of masses, the latter were relatively 
difficult to stain ; and, unless the viscid stuff had the limited 
physical course of fibres or networks, it was less easy than 
usual to kill, stain, or fix quickly the areas enclosed or cut off 
by it. 

In developing starfish and echinus eggs, there are rhythms 
in amount of time required for stains, or fixing reagents, to 
act, and these are sympathetic with rhythms of resistance 

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to pressure, of the whole eggs, or of the peripheral areas. The 
same held true of internal areas, which had rhythms of resist- 
ance, and at the same time showed like variation in resistance 
to pressure.^ 

If we simply throw eggs into some killing and hardening 
material, and wait for them to be dead enough, and hard 
enough, to resist the severe strain of subsequent operation> we 
necessarily miss much information from the substance as to 
how it is affected ; and how it reacts to the treatment. 

It seems to me such use of reagents should also be considered 
in the light of inducing pathological conditions; that the 
chemicals used should be watched as modifiers of the living 
substance^ — as drugs ^ in short. It is not impossible the science 
of medicine may thereby be indirectly aided. (See Selection of 

[25] Most remarkable of all physiological peculiarities cor- 
related with areas which are marked by stable or rhythmic 
viscosity of the substance, as associated with organization of 
its elements ; is an increased tenacity of life under adverse 

[26] I find that when an organism is crushed, or dissected, 
the power of continued existence is shown most specially by 
two groups of substance structures, whether the creature be 
Metazoan, or Protozoan : 

In the Protozoan, those portions whicb at all or most times 
maintain protoplastic activities, notably those which are lilose ; 
and those portions which show marked organization of the 
elements for contractile activities : 

Those portions of the Metazoan which at all or most times 
maintain protoplastic activities as their characteristic activity ; 
and those areas which show marked organization for contractile 

When an aquatic Metazoan is crushed, the muscular tissues 
are the last to die, — that is, to become vacuolated, and to 
have disorganization of the elements set in. 

The same is true of all aquatic Protozoa and the few endo- 
parasites I have examined. 

1 See also Activities ; — rhythms of viscosity in starfish eggs. 

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[27] And of all such structure in both classes of animals, 
modifications of the finer structure for contractile activity as 
in filose masses and pellicles, persist longest in an active state. 

Of higher forms, I have to offer as evidence in this connec- 
tion only such as is given by the crab and the frog. In the first 
class of structures cited, the protoplastic areas, the retention of 
primitive powers of local adaptation to environmental condi- 
tions, such as the lowest forms of the substance possess, which 
include a marked power of swift and stable organization of the 
elements, and of swiftly varying the viscosity of the interal- 
veolar stuff ; evidently serves to maintain in such areas a physio- 
logical existence, independent of the organism which in toto is 
the food provider for all areas. The limit of existence of such 
areas must be to great extent the limit of the stored up 
food, or stimuli, represented by their inclusions. 

In this connection it is hardly possible not to refer to the 
strange vitality and power of independent existence shown by 
those curious substance structures, the leucocytes of crab's blood. 
These, five hours after being drawn from the animal's body 
while it was in the soft-shelled state, still actively formed colonies, 
and exhibited free protoplastic phenomena. After general con- 
nection of the cells had been established throughout the whole 
by filose processes spun out to excessive delicacy, there was a 
marked effort on the part of those nearer the periphery to spread 
themselves out in quite a special way, in fiat, almost parallel 
processes, which again spread themselves out laterally, so as 
to form sheets and webs and mats of protoplasm ; producing 
at last a covering, or envelope, of very viscid substance for 
the whole. A good deal of migration of the individual cells 
was involved in these activities. The envelope had for the 
unaided eye a viscid, gelatine-like appearance, and when placed 
in strong light refracted from the surface. It was at the end 
of two hours so viscid and firm over the surface of a small 
wine-glassful of the blood as to sustain considerable pressure. 

The whole mass had now the value of a mass of protoplasm, 
or even an organism, for the filose processes everywhere bound 
the once single units into a firm union. The fluid surrounding 
them had the value, as to the external pellicle, of a mixed in- 

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elusion, interpenetrated by filose processes ; while the external 
layer had the value alike of ectosarc, and of a pellicle or pelli- 
cular membrane. 

Those leucocytes whose first position was nearer the center 
of the mass, behaved quite differently from those of the periph- 
ery. They coalesced changefuUy into ever-increasing lumps 
and masses of protoplasm, spinning threads and webs in all 
directions, even to the very periphery where they joined 
themselves to those units forming the membrane. 

There seemed to be shown here an organized effort on the 
part of members of the artificially formed colony to make an 
inclusion of the serum, and so to retain for the common sub- 
stance its accustomed environment as long as possible. (See 
Selection of Environment.) 

In the second class of structures, cited above as having per- 
sistent vitality ; the inclusions being already of an organized 
homogeneity in character, the power of the substance to vary 
local viscosity, and to maintain it, is thus emphasized, and 
necessary local food, or stimulus, is better conserved and con- 

In preparation for encysted states an envelope of increased 
viscosity is first formed ; sometimes there is merely increased 
viscosity of the pellicular membrane of the organism; some- 
times a separate membrane is formed by exudation, or possibly 
spinning phenomena, but this latter in these cases I have not 
seen. Echinus eggs were seen to form membranes by filose 
phenomena of the pellicular stuff. 

The breaking up of the original mass of a Protozoan into 
swarm-spores, or into macrospores, is possibly another mode of 
using a more viscous and finely organized state of the elements 
for preservation of the substance ; for these products are 
always surrounded by a very viscous and finely structured 
area, and, moreover, are supplied with motile appendages 
which are also tactile and prehensile, by whose means the 
substance seeks a new and more favorable environment for 

[28] I have observed in all young of Protozoa a markedly 
greater resistance to adverse environmental conditions ; a tem- 

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perature or state of the water which had caused death to the 
adult mass as such, seeming for these innocuous. Where 
death was escaped by the adult through formation of mac- 
rospores, these reorganizations of the adult mass showed an 
indifference to adverse conditions which was in many cases 
remarkable. I have seen Protozoa spores swim actively for 
many minutes in strong solutions of poison, such as corrosive 
sublimate, and to resist for some moments ammonia, the 
faintest trace of which actually disintegrated the adult. 

These facts may go to explain paitially the extraordinary 
resistance of bacteria and their spores to action of heat, cold, 
and other states inimical to the living substance. 

The prevalence of egg membranes and of ectosarc-like areas 
in embryonic states of Metazoan substance also receive here 
some light. 

[29] It was found that to shake starfish or echinus eggs, in 
early stages, not too hard, was productive of perceptibly greater 
resistance to reagents, to shaking and also to pressure ; in other 
words, the viscosity of the mass, or most likely of the peripheral 
portion, was increased by such treatment. The same result was 
had from pressure, not strong enough to rupture, between cover 
and slide. The substance here opposes an adverse environ- 
mental condition by a physical change of state. In this con- 
nection there are facts obtained in the study of contraction 
phenomena which are of vital importance. The reader is most 
particularly referred to the phenomena of cell-connection, both 
as seen in mechanical experiment and in the natural course of 

An excellent instance of rhythmic variation of viscosity is 
had in the pellicular membrane which forms the wall of the 
contractile vacuole of amoebae. This was found, contrary to 
the most widely accepted belief, to be not a stable, or per- 
sistent, substance structure. The subjects of observation were 
A. proteus and A, radiosa. 

At times in its rhythm, though not in each recurrence of 
this, the vacuole disappears ; not merely optically, but actually. 
For during collapse, which at some times seems to be more 

1 See Activities ; — rhythms of viscosity. 

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complete than at others, the pellicular substance becomes so 
completely relaxed, so fluid, that it mingles with the interal- 
veolar substance of the surrounding protoplasm, and the 
vesicles which at one time in the rhythm formed a more or 
less perfect alveolar layer about the swollen vacuole, are 
also mingled freely amongst those of the surrounding sub- 
stance, after the manner of endosarc. As the membrane 
re-forms after such collapse, its thickness, which at first is 
slight, is gradually augmented by access of interalveolar stuff ; 
and at the same time it undergoes more or less modification of 
its optical qualities, until near the most distended moments of 
the vacuole these qualities are at their maximum. That 
obliteration of the membrane is here a physical fact, and not 
mere optical illusion, is shown by the flux of the vesicles 
amongst each other, and also by the failure of even osmic 
fumes at such times to show a trace of substance modification, 
about the point of collapse ; while, on the other hand, during 
ordinary collapses, when the pellicle becomes merely plicated 
and difficult to detect optically, one can so preserve a very 
definite substance structure. 

Nor is the vacuole itself a permanent organ in the strictest 
sense, since, contrary to the accepted idea, it does not persist 
at one spot in the endosarc throughout the life history of the 
animal, but as I have witnessed, may fail after collapse to reap- 
pear where it has done so for hundreds of consecutive times. 
When this perhaps rare phenomenon occurs, the protoplasm of 
the immediate area has a very fluid, vague^ appearance, and I 
have seen the whole area, continuous substance as well as 
inclusions, excreted by a wave of contraction in the endosarc. 
Outside, it seemed to be a flocculent mass, which soon disinte- 
grated in the water and was dissipated. Later, after a consid- 
erable interval, another contractile vacuole was formed in the 
endosarc at another point, and the surrounding protoplasm of 
that region soon gained the more fluid appearance which 
usually characterizes the region of a contractile vacuole in the 
forms which gave these facts. 

It is no uncommon thing in Amoeba proteus and in other 
Protoplasta, notably the Heliozoa, to cast out thus portions of 

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their living substance, which seem to have become in some 
way weakened or effete. 

[30] The so-called membrane of the contractile vacuole is 
after all but an unstable, areal organization of the intend- 
veolar substance, as a contractile pellicle, or membrane, 
and besides its periodic efifacements, is subject at all times 
to the same flux of its substance as is the surrounding net- 
work. Yet in so far as it is at times so modified in its 
structure and reactions as to form, though but fleetingly, an 
area of organized physiological reaction ; and to function at 
such times as a contractile membrane; it is a true organ of 
the cell. 

It is a substance organ in the same sense as are all contrac- 
tile pellicles. 

To relate in detail all the phenomena observed in the con- 
tractile vacuole of Amoeba proteus alone, the rhythmical sub- 
stance and structure changes, would fill a large essay, which it 
is hoped to edit separately at some future date. 

In some Protoplasta, the number of contractile vacuoles is 
not fixed, new ones arising capriciously in their midst. This is 
true of some Ciliata also. 

I find that the contractile vacuole of Protoplasta is, as 
a rule, but not always, discharged to the exterior of the 
mass. At times, collapse takes place before the periphery 
is reached. 

A strange intermittent contraction is found in the pellicle 
which lines the pharyngeal and oesophageal cleft in the Vorti- 
cellidae. By an annular constriction, the lower end of this 
cleft which runs from the peristome down to the lower end of 
the body is cut off, and then with its contents forms a sharp- 
ended, pear-shaped sac full of water and of ingested food. 
The contraction by which it was cut off passes on behind this 
sac, pushing it further and further from the end of the now 
much-shortened tube. By continuance of this same contraction 
in a wave-like impulse, the sac is forced upwards through the 
endosarc, along a gracefully curved course, until by pause of 
the contraction it rests at some point. During this time, by 
relaxation of the pellicular substance, it has yielded still more 

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fluidly to the laws of its structure, the foam structure, and 
becomes a spherical food sac.^ 

Here we have a physical pellicle of the substance, formed at 
moment of a splitting apart of the endosarc, modified into a 
true contractile pellicle. After functioning for a time in this 
way, it is as a viscid fluid transferred to another portion of the 
endosarc, and there being for a time relaxed, functions merely 
passively and permits the passage of fluid through it. Then 
it may more or less often function again as a contractile 
membrane during the digestive process. Finally, it is again 
forced outwards by a contraction wave of the surrounding 
substance and beneath the peristome collapses, expelling its 
contents to the exterior, after the manner of a contractile 
vacuole ; or it may itself also be expelled by the endosarc and 
perish as waste matter with its contents. 

Beside the formation of digestive sacs, these activities of the 
substance were seen to be made use of to rid the organism 
of unwelcome presence in the pharyngeal cleft of large num- 
bers of bacteria which were whirled into it by the cilia. 
Under similar circumstances many rotifers would simply 
turn their digestive tract inside out, and so get rid of the 

Here, with no visible nervous organization, the same result 
was had in rather a more complicated way ; for the end of the 
pharynx, after being pinched off, was forced all the way up 
through the body mass and expelled, walls and contents at 
once into the water. 

[31] In a single life phenomenon there is here shown a 
selective power of the substance ; a migratory power as a sub- 
stance organ ; metamorphosis of both substance structure and 
substance function ; rhythmic changes of viscosity ; and above 
all there is shown a correlation of impulse, a coordination of 
these activities throughout the entire mass, which is as perfect 
as that seen in organizations, where, to make it more com- 
prehensible Q)y there are enormously complex systems of struc- 
tural modifications of the substance. 

^ These appearances were construed by Ehrenberg into a complex oesophageal 

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To sum up all that has here been brought forward as to 

[32] Pellicles are of the interalveolar substance; and we 
shall presently see, that here, as elsewhere, the substance 
preserves in toto its inherent powers; so that there are not 
found in masses activities and characters, which are not 
found also in pellicles and small areas of the interalveolar 
stuff — areas so small that we cannot assign a limit to their 

Pellicles are, characterized by a hyaline, and, with all but the 
highest powers, a structureless appearance. Their inclusions 
are for the most part fluid ; if we exclude the so-called " gran- 
ules," they are optically wholly so. 

(a) Pellicles, whether as passive masses of interalveolar 
foam, or as contractile membranes, are organizations of the 
two groups of elements of protoplasm, on a basis of its visco- 
fluid foam character. 

(b) This organization is in direct correlation with physio- 
logical activities of the substance ; and hence I hold pellicles 
to be true substance organs. 

(c) They represent the necessary primitive type of physio- 
logical organization of the substance ; and are therefore typical 
of the primitive type of substance organ. 

(d) They epitomize also all typical substance organization 
for reaction, in character, to environment. 

(e) They are the primitive, as they are still the characteristic, 
end-organ of the living substance, as such. 

(f) They have, therefore, in this respect also, the character 
of all true ectosarcal formation. 

(g) Pellicles are the primitive ectosarcal formation, and 
are, in their origin as well as function, typical of all ectosarcal 

(h) Because of both their form and function it seems to 
me impossible to deny to pellicles the value of membranes, of 
living membranes. For the substance as such, they are mem- 
branes par excellence. 

There is the more reason for this view since all those struc- 

^ See Activities, — also Substance as Such. 

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tures to which is conceded the value of a membrane arise as a 
simple or complex pellicular membrane.^ 

[33] In the pellicles of artificial froths, osmosis is controlled 
by the passive physical conditions. In living foams, osmosis is 
controlled also by changes in the pellicular substance, which 
are correlated with a physiologfical function or power of the 
substance (contraction). This is proven by the fact that exist- 
ing physical conditions are in a moment set aside under such 
diverse stimuli as are supplied by reagents, or even mere 
pressure or shaking ; the visible phenomena in all cases coin- 
ciding with those correlated with contractile function of the 
substance in typically contractile areas. 


[34] Although in living protoplasm a true "alveolar layer" of 
Biitschli may very often be found beneath the pellicles which 
limit masses, and also those which surround inclusions, as the 
contractile vacuole, these appearances are in most cases transi- 
tory or transitional only, and are constantly disturbed by the 
action of other forces. Excepting cuticular and other substance 
structures which have organized contractile activity, — and 
these may occur at any point in a mass, — so-called " alveolar 
layers" among the Protozoa and among many Metazoan struc- 
tures are subject to constant disturbance. 

(a) On the other hand, in dead or dying protoplasm, in some 
peculiarly relaxed, inert, or partially atrophied areas and masses, 
in short, wherever and whenever there exists a marked relaxa- 
tion of the substance, alveolar layers are beautifully distinct, 
and form at points in relation to the mass which also bear out 
their physical origin. 

It is significant that in almost all preserved material, espe- 
cially where the method has been rather slow and sufficiently 

1 Membrane (Century Dictionary definition) : A thin, pliable, expansive struc- 
ture of the body ; an expansion of soft tissue, or part, in the form of a sheet or layer, 
investing or lining some other structure, or connecting two or more structures. The 
term is used in the widest sense with little or no reference to the kind of tissue 
which may be involved, the membranous quality depending upon thinness and 
pliability, not on texture or fabric 

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delicate to avoid gross vacuolation of the structure of Biitschli, 
alveolar layers are marked. 

The fact that in the living substance these layers are less 
stably present than Butschli supposed does not militate against 
his theory of the structure of protoplasm. It is, on the con- 
trary, enforced, because the moment the substance as such 
is released from certain physiological controls, it proves its 
physical form by acting as a mere visco-fluid foam. 

[35] While I find alveolar layers to be expressive of the 
physical form of the protoplasmic foam, they are not neces- 
sarily due to this alone, but may express also effects of physio- 
logical states and physiological activities. 

Many alveolar layers given by preservation methods express an 
osmotic vacuolation of the living structure. This can be proven 
by use of reagents on living material under the microscope. 

[36] Contrary to all the conditions found necessary by 
Butschli for alveolar layers in artificial foams, I find the living 
substance forming freely, in a moment, at any point in its mass, 
and while the general structure of Butschli is of markedly 
irregular size, and the mass or area is of marked viscosity; 
perfect alveolar layers. 

The most interesting instance I know of this is afforded by 
the formation in developing echinus eggs of a perfect and 
double alveolar layer internally to the cell-mass. This after- 
wards becomes for a short time part of the external peripheral 
layer of the new cells, the split taking place between the two 
rows of vesicles. 

[37] Marked viscidity of a mass, or area, of protoplasm 
seems no bar to formation of a very perfect alveolar layer ; the 
irregularity of its existing structure, no impediment ; nor, on 
the other hand, is the fluidity of a living area or mass any 
guarantee that there we shall find a definite alveolar layer, 
except more or less unstably. The living foam, then, is able 
in this respect to defy, or set aside, those controls which rule 
the purely artificial foams of Butschli. 

In artificial foams the finer vesicles tend to range themselves 
peripherally to the mass, the outermost layer, in foams which 
are not very viscid, being the alveolar layer. 

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[38] I find in living protoplasm that the size of the vesicles 
of Biitschli's structure cannot be brought under any rule cor- 
relative with their position in masses. 

In any portion they may be large or small, or they may be 
either in turn, and their organized arrangement extend in any 

Except the fact that the pellicular formations are always of 
the finer foam of the interalveolar stuff, it is impossible to 
group in point of size the remarkably diverse structural facts 
of protoplasm. 

The same results are attained by the same creature, within 
short intervals of time, by widely different distributions of the 
substance elements. Peripheral areas may be coarsely or finely 
vesiculate. They may be very viscous and have a finely sub- 
divided structure, and then one may group together numberless 
instances of similar coincidence. They may be very viscous and 
have a very coarse alveolar structure ; and here again one may 
summon many like cases. They may be markedly fluid and 
have either coarse or fine structure. A coarsely structured 
area may at one moment be very fluid and at another very 
viscid. The same is true of an area or mass having marked 
viscosity. In such forms as Amceba radiosa the substance 
may be watched for days varying from moment to moment its 
play amongst such phenomena. 

These changes of physical state are most instructive, for 
they represent the characteristic powers and activities of the 
living substance. 


For substance differentiations, as found in general researches 
among many, more or less scattered, organisms, it is difficult to 
lay down general rules. It might seem well-nigh impossible, 
so contradictory are the evidences when followed along some 
lines. Yet it is hard to work long on the substance as such 
without becoming sure that there are generalizations to be 
made, and hardly to be escaped, since along other lines the 
habit of the substance is clearly defined. 

[39] Thus much is certain, that wherever was found organized 

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physiological activity, there was found also physical organiza- 
tion of the two groups of elements of Butschli's, or a finer, 
or a coarser, foam structure, with or without further physical 
modification of the gfiven interalveolar structure. As to the 
observer's perception, this organization arises sometimes simul- 
taneously with the activity, sometimes precedes it, and may 
outlast it, but always accompanies it. 

(a) It does not necessarily follow that the structure visible 
to the observer in a physiological area is that which is to be 
correlated with the gfiven function ; nevertheless it is true, 
as will be shown, that function and vesicular structure are 
indeed too closely linked for us now to set them apart. 

(b) Capricious as seems the substance, unstable and incal- 
culable as are its manifestations, there can yet be found certain 
broad laws, or rather habits, of structural arrangement under 
which it is wont to express itself. 

(c) Under the head of pellicles somewhat was indicated of 
the manner in which masses of protoplasm are subdivided and 
differentiated structurally by arrangement of the continuous 
substance and the inclusions, and by the character of these two 
sets of elements. And to pellicular formation much of the 
optical, as well as physical and physiological, differentiation of 
areas is due. (See also Ectosarc.) 

The majority of facts relating to these things are so inextri- 
cably correlated with phenomena of the substance as such, 
which must be grouped under the head of activities, that they 
will be treated of in that section, but some few important gen- 
eralizations are offered here. 

[40] Two groups of areal differentiation are most marked, 
— those in which the function is of a vegetative sort, and those 
in which it is organized or emphasized irritable or contractile 
function, or both. 

In the former case the continuous substance has not usually 
that marked refractive and viscous emphasis, stable or rhythmic, 
which peculiarly characterizes the other group of structures, 
yet such there may be, at intervals during active secretion, as 
was seen in the gland of a certain rotifer {Megalotrocha)^ 
which, situated by the point of exit, pours out a glairy sub- 

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Stance about the eggs as they leave the body.^ There are cases 
in which the two sorts of structure are combined. In these 
the substance, both as such and as area or organ, functions for 

In areas having marked glandular function for the organism, 
the inclusions corresponding to Biitschli's structure are apt to 
be much enlarged. It is noteworthy that certain brain and 
ganglionic areas of some rotifers showed such a state of things. 

Areas whose function is not for displacement but in gross 
chiefly of a chemical nature, are thus of the vegetative class. 
Hence all areas where the substance as such occupies itself 
chiefly in digestion or assimilation processes join this tentative 

[41] Wherever areal function is markedly irritable or con- 
tractile, the inclusions of the organized substance are uniformly 
fluid, and the discontinuous element is evenly distributed with 
respect to the course of organized action of the continuous 
element.* It is evident that the elastic, contractile, continuous 
substance has freest physical opportunity when its inclusions are 
fluid and uniform in size, 

(a) Further, whenever in areas characterized by heterogeneity 
of inclusions there arise organized activities of this sort, how- 
ever fleeting, a precisely similar state of things is associated 
with them. In development of many eggs the direct action of 
the substance with respect to its different inclusions is striking. 
Solid substances, such as yolk or pigment, are segregated or 
deported temporarily from areas in which activities of the sort 
under discussion are to appear. Sometimes these areas are in 
relation to individual cells, sometimes in relation to the whole 
coming organism as a substance mass. 

[42] Wherever, in short, the substance is preparing itself for 
organized contractility or irritability (if, indeed, we may so far 
separate the two), it provides for itself just those physical con- 
ditions which offer greatest vantage.* 

^ In this form I find there is a special opening for the egg to pass through, 
quite distinct from the cloacal opening, and situated just above it 

* See Striation ; also below, nervous areas. 

' Obviously this might be conversely stated* yet, it seems to me, with less 
justification as yet. 

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[43] The most readily noted feature of such organization is 
uniformity in kind, size, and arrangement of inclusions. But 
this is in most cases secondary in importance to differences 
in quality, structure, and position of the continuous substance. 
These things are as true of contractile and transmissive areas 
in the Metazoa as in the Protozoa ; and portions of one or 
many "cells," may be involved in these modifications. 

(a) In areas formed for interaction with environment, whether 
limited to pellicles or involving an underlying alveolation, fluid 
inclusions are gathered together, arranged with great uniformity, 
and often subdivided to a point of gi'eat minuteness, which 
latter change involves, of course, a physical extension of the 
lamellar substance. (For the full significance of these things 
the reader must see the later sections.) 

(b) Such are ectosarcal formations, and such the arrangement 
of the elements wherever the substance meets internal contacts 
with organized reactions characteristic of ectosarc. 

(c) All areas to be grouped with ectosarc, that is, all areas 
which are formed of interalveolar material, with or without 
fluid inclusions of the structure of Biitschli, have the same 

[44] There is a marked habit of the substance to meet all 
contacts involving a number of the alveoli of a given structure, 
with organization of its elements in such a way as to produce 
an ectosarc-like formation. This may be formed of inter- 
alveolar material alone as a pellicular modifier, or may involve 
more or less of the structure of Biitschli. 

(a) In the formation of ectosarc there is a tendency to reduce 
the structure by subdividing the alveolar inclusions. 

(b) There is a tendency, too, in such areas, whether so sub- 
divided structurally or not, to increase of viscosity. 

[45] I find that true optical homogeneousness, or structure- 
lessness of protoplasm, is most often produced by minuteness of 
structure, rather than by mere stretching of alveolar lamellae. 
The latter state produces, of course, a less dense and refrac- 
tive substance, while the former produces a more dense and 
refractive substance. So-called, as well as veritable, optically 
structureless protoplasm is commonly denser, more viscous, and 

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markedly more refractive than the structured areas. While I 
agree with Biitschli that we are often deceived, as in the in- 
stance he cites, and which I have also verified, into believing 
an area of protoplasm to be structureless because of a greater 
tenuity, by stretching, of its alveolar lamellae ; I have seen 
numberless times the origin from visibly structured areas, of 
true, optically structureless protoplasm, by progressive sub- 
division, or reduction, of the structure of Biitschli. This often 
takes place with marvellous rapidity, and to a point of minute- 
ness causing great difference of optical quality, and the area so 
formed may be very fluid or markedly viscous. 

It is in no other way that the ordinary structureless portions 
of ectosarc of Amoeba, etc., are formed. True, there are very 
hyaline and delicately structured areas which arise without 
such reduction, except near the periphery, by outflow from the 
endosarc of interalveolar material, with or without fluid alveoli 
of Biitschli's structure, but it is reduction phenomena which 
give a veritable structurelessness of appearance. Both sets of 
phenomena may be witnessed within a few moments in an 
Amceba radiosa which delights in varying its activities a thou- 
sandfold. Even without any perceptible change of form in the 
animal, which may at the time be somewhat rigidly extended 
in the water in the form of a star, of four, six, or any moderate 
number of arm-like rays, or spread out as a fan-shaped film ; 
the structureless substance will in a variable time be resolved 
again to a very marked structure of Biitschli. 

At both times, under sufficiently low powers, the ectosarc will 
have an optical structurelessness, and possibly also a greater 
refraction than usual ; yet in one case the structure of Biitschli 
as such has been obliterated, and in the other case is again 
present. The increase of refraction in the substance may be 
seen to be caused by a change in quality of the interalveo- 
lar material, so that the meshwork of Biitschli's structure stands 
out refractively, as thickened trabeculae, under lower powers 
than are usually needed to resolve it. Structural reduction in 
the living substance does not always result in thinning the 
interalveolar material. Pellicular and interalveolar material 
furnish commonest instances of apparent optical structureless- 

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ness, yet in them a true structure has been found, and one under- 
going such modifications as are found in the coarser structures. 
It is necessary to distinguish between visibility of areas and 
visibility of the protoplasm proper, or continuous substance, for 
the character of its inclusions may make a whole area distinct 
or prominent, while its protoplasm proper may at the same time 
be very fluid and not so readily seen as that of adjacent areas 
which in toto are less conspicuous ; but the optical phenomena 
in such cases are markedly different from those in which areal 
visibility is due to quality of the continuous substance. 

[46] I would group all areas having the origin and character 
of the ectosarc of protoplasts under the head of ectosarcal 
formations or modifications of the substance. As stated above, 
all physiological pellicles fall naturally into this group. 

To glance for a moment at the formation of ectosarc by 
Amoeba may enable the reader better to understand the argu- 
ment. The point involved is of radical importance. An 
Amoeba proteus is at all times covered by a variably thick, 
hyaline layer, whose surface forms an unstable pellicular mem- 
brane. But when at times, in the heave and flow and evanescent 
shiftings of the protoplasm, there is at any point a rush of sub- 
stance that wholly or partially pushes, or breaks through, this 
layer or is yielded to by it, so as to come in more or less close 
proximity to the water, the phenomena may be said to represent 
to some extent a primitive formation of ectosarc. This may 
have been primitively a purely physical incident, taken advan- 
tage of by, or necessarily causing physiological or physical 
vantage for, the living substance.^ 

One does not, however, see the whole mass covered at any 
one moment by such an area ; that has been the work of ages, 
and the present habit, while typifying a more primitive act, is 
of course a much modified and extended form of it, for in the 
simplest Amoeba known, the plastic substance is already very 
old and full of regfistered experience. 

Watching an initial formation, not a mere displacement, of 
ectosarc, one sees a gradual, or more or less sudden, or even a 
spurting, outflow of what has been called the hyaloplasm of the 

^ See New Structural Formula. 

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endosarCi that is, the more hyaline, fluid, colorless portion which 
does not carry solid bodies or nutritive material in an unassimi- 
lated state. This is already, in esse as well as in posse^ an 
ectosarcal formation, and may be seen to function in a charac- 
teristic, organized way as it is mingled with the true endosarc. 
Besides that hyaloplasm which is formed of the structure of 
Biitschli, there is an interalveolar, hyaline substance, the finer 
foam, which also flows out with, or besides, this, and the two 
structures mingle fluidly amongst each other. In such ecto- 
sarcal outflow, there follows the first impulse a more or less 
active motion of these alveoli amongst each other. There are 
so many modifications in individual cases of the result in 
particular, that it is difiicult to seem to describe accurately 
any one instance, unless to those who hear all are familiar. 

The damming back of outflowing substance by a rapidly 
effected pellicular formation of ectosarcal character, or by 
obstruction of already existing pellicle or ectosarc ; as conflict- 
ing with the onward rush of material from the endosarc, 
causes often in the newly formed ectosarc a kind of boiling 
motion which comes to an end variably soon according to 
circumstances. Where the barrier is an existing ectosarcal 
formation which does not readily yield to the outflowing sub- 
stance, the direct course of this is changed to more or less 
lateral flow. The alveoli of Biitschli's structure then become 
less freely mingled with one another, as if the impelling force 
ceased to act, or as if there arose an increased viscosity crossing 
the path of the latter, or sometimes following it in point of time 
and impeding the fluid motion. Such viscosity is understood 
from certain optical changes in the quality and action of the 
substance, or by changes in structure. There is little doubt that 
further progress is often checked by formation also of a more 
finely structured, and therefore more viscid and resistant, area 
in the advancing substance itself, along the line of its contact 
with the existing ectosarc ; just as in this latter when it came 
in contact with the water, a similar area was formed, checking 
in the same way the substance following fast behind. 

If the outflow of hyaloplasm has burst through an existing 
pellicular membrane into the water, or runs very close to this, 

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there is more apt to occur this phenomenon which is very 
characteristic of ectosarcal formation, and which has almost un- 
bounded possibilities in an evolution of protoplasmic structures, 
as will be shown further on,^ and in still another publication 
now in preparation for press. I speak of what I shall call 
structural reduction, that is, a redistribution of the elements in 
an existing structure so as to group them into a new and finer 
foam structure. The vesicles of the existing structure become 
quickly more minute, and should subdivision continue, yet finer 
and finer, being always increasingly small towards that outer 
side or portion of the area which is peripheral as to the mass. 
Structural reduction takes place in a number of ways. The 
alveoli of Biitschli's structure were seen at times to be cut in 
two by constriction, or pinching off, just as a contractile vacuole, 
or as the oesophageal cleft of Vorticellidae divides. Sometimes 
the interalveolar substance spins itself into and across fluid 
inclusions ; sometimes there seems to be local relaxation of 
interalveolar or continuous substance, causing redistribution 
of alveolar inclusions. 

But however it may be done, the fact is certain that it is 
done, and that in most cases of ectosarcal formation there is 
more or less structural reduction and reorganization of the two 
groups of elements, which may or may not extend itself to the 
interalveolar structure where it is visible by change of the 
optical quality of this substance ^ or again may be confined to 
this alone. 

The amount of such reduction varies with the circumstances 
of formation, or even, to all seeming, irrelevantly to these. 
Where large areas of new ectosarc are brought into direct 
peripheral contact with the water, or general environment of 
the mass, there is more structural reduction than where the 
outflow is checked by presence of existing ectosarc in a thick 
layer. It was the optical appearance of these phenomena as 
seen with powers too low to resolve structure, which gave earlier 
observers an idea that the substance became coagulated on 
contact with the water. As a matter of fact, ectosarc may be 

^ See Selection of Environment. 
* See Striation. 

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far more fluid than endosarcal areas, although it usually is seen 
in a more viscous state. 

One interesting unstable area of structural reduction which 
I think has not yet been described, was seen as a variably 
present and fluctuating area surrounding the nucleus in 
Amoebae. Very finely vesiculate, often passing into true 
optical structurelessness, when it had an effect of brownish or 
yellowish hue and was of a dense and peculiar appearance, this 
viscid mass adhered to the nucleus and was carried along with 
it, as the endosarc rolled hither and thither. Less transparent 
and also far less refractive, though quite as dense nearest the 
nucleus, it reminded one much of the <' achromatic ** areas in 
embryonic masses. 

Its amount was greatly varied from minute to minute, some- 
times being at least one-sixth the width of the cytoplasmic 
area, at others merely a sticky-looking stratum close to the 
nucleus, as if the cytoplasmic pellicle were unevenly aug- 
mented. Whatever its thickness, it swept along with the 
nucleus, holding its place as an almost solidly attached sub- 
stance while its outer portions were drawn out as threads and 
viscid-looking processes in the cytoplasmic, flow over its sur- 
face, 80 that it looked somewhat like a caged Heliozoan in 
closest physical union with the surrounding fluid alveolation. 
No special life phenomenon was seen to be correlated with the 
presence of this area. 

It is not only at time of initial formation of ectosarc that the 
phenomenon of structural reduction is seen. In ectosarc long 
formed and quite stably persistent through long periods, and 
in ectosarcal organs and areas, similar redistribution of the 
elements takes place at times, changing the physical form of 
the substance locally or throughout the whole area from a 
perhaps marked and uniform structure of Biitschli, or finer 
forms, to others progressively finer until a point of actual 
optical structurelessness is reached. From this state redistri- 
butions may later bring before us again a true structure of 
Biitschli, not the former one but another, in which it is more 
than likely not one particle of the elements holds its initial 
relative position as to the others or as to the whole mass. 

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It is important to remember that in the living substance true 
ectosarcal layers are formed at any depth and in any relative 
position and of any form, as readily as they arise externally to 
masses ; and that such formation may be the work of a moment 

[47] Ectosarcal formations often arise by outflow of the 
interalveolar substance alone without implicating or even visibly 
disturbing an existing structure of Biitschli.^ 

[48] All ectosarcal formations appear to be essentially organ- 
izations of the continuous element involving redistributions of 
the alveolar inclusions ; and to be formed for physiological func- 
tion ; for organized reaction of the substance as such in its own 
character to environment. 

[49] In many areas so formed it seems to be of no moment 
which portion of the continuous element is in contact with 
any visible vesicle, but all portions can react alike near all 
portions of the general inclusion fluid. In other words, there 
is in such areas a homogeneousness of the continuous element 
and equal homogeneousness in general character of its spe- 
cific stimuli. Any part of such areas is, therefore, for the 
time being physiologically equal to any other part in these 

(a) Yet any part of the area may show specific differences 
of action, which are secondarily referable to either local dififer- 
ences in its own finer structure, or inclusions, or slight 
differences in kind of the inclusions of the given structure. 
Such local specific activities are always directly referable to 
characteristic changes of the continuous element. 

[50] Although true protoplastic ectosarc, as it is best known, 
is commonly seen as arising in response to contact with external 
environment, it may hardly be thought that such areas are in 
general a result of such contact, for in the majority of cases 
they are formed before, often long before, the occasion of their 
use. Sometimes they may never even reach that environment 
for reaction to which they have been moulded. In the forma- 
tion internally to a Protozoan's mass of pellicles and pellicular 
organs which, till after completion, are shut out from their 

^ See Activities. 

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specific contact as organs with external environment, are seen 
instances of areal differentiation on an ectosarcal basis preceding 
physiological function : of vesicular preformation of substance 
organs before the actual opportunity for their use has arisen. 

Among the Metazoa such redistributions are common and 
numerous in development of cells and organs. 

Within the endosarc of Amoebae and other Protozoans one 
can see, fleetingly formed from moment to moment, small local 
areas of true ectosarcal character ; that is, showing a uniform 
distribution of the two sets of elements in relation to each 
other, with homogeneousness of both in general character ; and 
with certain organized, though fleeting, activities which are 
characteristic of true ectosarc. Besides these, there are the 
stable pellicular formations about nucleus and food sacs and the 
contractile vesicle, which are all organized areas of interalveolar 
stuff involving more or less of the structure of Biitschli. Like 
other contact areas, or substance organs, they are all charac- 
terized by fluidity and uniformity of their inclusions. 

Such true and stable alveolar areas as are found in Protozoa 
are, as a rule, marked by organized contractility, so that these 
also are less expressive of physical form than of the correlation 
with the foam structure of certain intrinsic powers of the 

With few exceptions, ectosarcal formations show with vari- 
able intermission contractile activities. Even the nuclear 
pellicle cannot be excepted, for few nuclei do not from time to 
time have amoeboid changes of contour more or less marked. 
^Wherever in a Protozoan or Metazoan contractile activities 
were visible as such, there was an organization of the elements 
and structure typically ectosarcal, no matter how fleeting the 
physiological manifestation might be. The structure might be 
stable, or might not outlive the activity, but in all cases organ- 
ization of the continuous substance upon a fluid-inclusion basis 
existed at the moment of function in the functioning substance, 
and was exactly of the sort seen in typical ectosarcal areas. 
Wherever preparation of areas for organized contractile func- 
tion was watched, the process was seen to be not other than 
that typical of the formation of ectosarc. 

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Whenever organized contractility appeared for the first time 
in any existing, or new, irregular ectosarc, it was preceded by 
thorough and systematic reorganization of the elements on the 
same basis as that of the initial formation; the interalveolar 
substance having always the leading r61e. 

And wherever such a characteristic arrangement .of the ele- 
ments* already existed, organized contractile activities were 
seen to arise at some moment without further change in the 
existing structure than such as pertained to that rhythmic 
qualification itself. 

[51] To sum up: contraction was found always associated 
with an organization of the elements which is typically ecto- 
sarcal ; and typically ectosarcal areas were found to be charac- 
teristically contractile. Organized contractility was not found 
except where the elements either of Biitschli's or of the inter- 
alveolar structure were directly or indirectly shown to have 
such a constitution. 

It may seem, perhaps, that the same things have been said 
a great many times over in slightly different ways. This is 
exactly what nature does with these facts ; and it is only by 
following constantly along different, yet closely kindred, lines 
up to the same conclusion that one gets a realization of the 
truth in this matter ; a fundamental truth it is with respect to 
the living substance and its habits of self-expression, and there 
is none other, I believe, so important in evolution. 

[52] In the light of such facts, ectosarc appears to be the 
type, or illustration, of all characteristic substance organization; 
for, as it will be shown that contractility is still the basic, ^ 
typical activity of the living substance, as far as this can be 
seen to act, so also down to this limit is ectosarc formation a 
fixed, physical basis for this activity; the basic mode therefore 
of organization of the characteristic powers of the living sub- 
stance. That this relation is indeed inevitable from the nature 
and relations of the powers and the form of the material, will 
be shown in the section on contractility and in that on the new 
formula offered for the living substance. 

It has been stated that the stability of a structure of Biitschli 
is no guarantee of stability of the living substance as such in 

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that area. This is true whether the area is one stably organized 
for contractile activities or vegetative in its habit.^ Even in 
strongly contracted masses and areas having a marked struc- 
ture of Butschli there are numberless local disturbances of the 
interalveolar substance, even transposition of this which is yet 
responsible for the structure and fxmctional activity of the area 
in gross. After organized function throughout the area, por- 
tions of the contracted substance relax, but the characteristic 
organization of the structure remains marked and stable in the 
whole area. This shows certainly that not all of the continu- 
ous substance is necessarily involved in maintaining contrac- 
tion of any area, on the basis of a given structure. 

In the few Metazoan organisms whose development I have 
watched, there is a more or less gradual formation, peripheral 
to the mass, of an area which can be grouped with typical ecto- 
sarc alone. And for each so called cell mass, or nucleated and 
pellicularly separated subdivision of the general mass, there is 
also such a layer.' As the Metazoan substance, in its life- 
rhythm as organism, approaches an adult stage, there gradually 
comes to be by successive reorganizations of its elements, an 
area of true ectosarcal type in gross. This has been called the 
ectoderm, and is formed of "cells," to the general substance of 
each of which the same typical structure may be extended in a 
variety of ways. Of such Metazoan ectosarc are made, as a 
rule, all areas of the mass, or organism, which as organs corre- 
spond in function with true ectosarc in Protozoa; that is, which 
act as intermediators between the mass and its external environ- 
ment. True ectosarcal formations of the Metazoan are, how- 
ever, by no means confined to such cells; but most, or perhaps 
it may even be said, all other cells form to some extent such 
areas of their substance.^ Nor can these be held identical in 
any way with that purely physical ectosarc of artificial foams, 
since in their origin the living substance seems to be neither 
dominated wholly, nor inhibited, by those physical conditions 
which Butschli has shown us do, in artificial foams, produce, 
or forbid production of, such areas. 

^ See ActiTities — filose ; also Contractility. 

* See Activities, starfish and sea-urchin development. * See Striation. 

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[53] Ectoderm with its products is but either repetition in 
gross, or a multiplication by repetition, of causes and effects 
which form the Protozoan ectosarc and its products. Beneath 
the seeming demarcation of "germ layers" with their pro- 
ducts, the substance expresses itself again, more definitely, in 
a series of minute but kindred structural organizations of the 
elements. These are to the substance as such what the germ 
layers are to the substance as organism. There are through- 
out the whole Metazoan organism countless areas of wide 
range in size; some restricted to the limit of single cells, 
others passing through large numbers of these, some of most 
minute vesicular structure, some having a beautifully marked 
structure of Biitschli, or even coarser vesiculation ; but all 
taking their rise from the general or "undifferentiated" proto- 
plasm ; all functioning in a kindred way towards substance 
environment ; and all true ectosarcal formations in origin^ 
structure, and activities. The same is true of endosarcal 
areas, for these are found in strictly ectodermal regions and 
cells of the organism. 

Physical nature heals the wounds of the living substance 
with a frail pellicle of continuous substance ; physiological 
nature supplements, or supplants, this as rapidly as may be with 
a formation of ectosarc. In simpler and more primitive types 
of organism this service is performed by interalveolar substance, 
often aided secondarily by areas of Butschli's structure. In 
more stably and complexly organized, cellular types the same 
surgical office is given by protoplastic activities of neighboring 
cells; by wandering cells, as leucocytes, or migrating cells, and 
as far as possible the damage is thus primarily repaired. 

By so much of experiment as was made, it is certain that 
living protoplasm tends to protect itself, directly or indirectly, 
by various devices from unsympathetic environment, albeit of 
a kind never before experienced; and to protect its internal 
environment from change thereby. Such devices are of a 
nature which seals the substance for a time from intrusion of 
adverse or disturbing conditions, and bar it in with its own 
existing and intrinsic conditions acquired under more favorable 
auspices. That is, the living substance shuts itself up with its 

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own internal resources, its hoarded specific environment, so 
that for a variable time it becomes wholly independent of 
external environment, of even the element it usually respires. 

From the formation of ectosarcal areas as external protection, 
to the natural habit of encysting, so common among the Protozoa 
and retained by protoplasm even among such highly differen- 
tiated forms as rotifers, to the self-sealing habit of snails, etc., 
and beyond these to the habits of hibernating found among 
the higher animals, even mammalia ; the phenomena are strictly 
to be grouped together under this interpretation.^ 

Through a wide range of forms, protoplasm makes use of 
ectosarcal layers for purposes of temporary isolation to a more 
or less complete degree from the direct influence of external 
environment. The purpose subserved may be an economy of 
energy, and of material both as hoarded reserve and as a factor 
of immediate environment during metamorphic changes ; or it 
mjfy be purely protective during adverse conditions of environ^ 
ment. A very common thing is for part of the organism, or 
all of it with small local exemptions, to be superficially pro- 
tected from external environment by such formations more or 
less modified secondarily ; and such is the origin of all exo- 
skeletons, and indeed of many endo-skeletons also. 

The lower forms, and also the higher forms in embryonic 
stages, make large use of ectosarcal layers produced from their 
mass so as to form distinct membranes which either produce 
substance for a non-living membrane or themselves atrophy 
later and are discarded after the need for seclusion is ended. 
The striated membrane in Echinus eggs is of this sort. I 
have watched the progressive formation of silicious capsules in 
a fresh-water Radiolarian, Clathrulina elegans^ upon a basis of 
a coarsely vacuolate peripheral layer of ectosarc, and noted an 
ectosarcal origin for the stalk ; I have watched also the gradual 
formation, by progressive rearrangement of the elements with 
subsequent chitinous transformation, of the very complex 
capsule of the winter egg of the fresh-water Polyzoan, Crista^ 

These phenomena are one with the partial independence 

^ See Selection of Environment ; also Fosterhood. 

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and physiological isolation gained for di£Ferent areas or sub- 
stance organs by pellicular formations of ectosarc. When looked 
at from the standpoint of protoplasmic habit, they must be 
regarded as continuous with the protection by ever increas- 
ingly complex and complete ectosarcal areas of all portions of 
the organism not concerned in the ingestion of food ; and finally, 
with the systems of protective apparatus which, as tactile, and 
as o£Fensive and defensive structures, guard at last even those 
weaker regions. The original formation may undergo more 
or less subsequent modification by secretion, or deposit, within 
or about it, of non-living material, such as chitin, silica, or 
the like. The actual living substance may be withdrawn, or 
may be gradually atrophied. In many cases it is impossible 
to say whether it is still present at a given stage, or whether 
it has been withdrawn or atrophied. 

The function of secretion does not inhibit in the substance, 
nor imply its renunciation of, typical contractile characters ; 
on the contrary, the latter may accompany the former to a 
marked degree. There is even strong reason for supposing 
that the function of contractility is always correlated with more 
or less excretion from the functioning substance, and here again 
it must be urged that it is most difficult for us to draw a line 
as yet between substance secretions and substance excretions. 
* For we are dealing always, it must be remembered, with a 
complex series of alveolar structures, not with one alone, as 
Biitschli's, for instance. Rhythmic, or irritably responsive, 
contraction most frequently accompanies a secretive function, 
and typical ectosarcal characters are very favorable to this 
state of things. By contraction, interalveolar vesicles may 
readily be emptied into the larger inclusions of Biitschli, and 
from these, which are storehouses of secretions as we know 
them, contractions of their interalveolar material pour the 
fluids. In strict sense, though not in common usage, the sub- 
stance produced by areas and organs should be called excretions 
when exuded from those places of formation, and secretions so 
long as they are considered in situ, although here they may be 
again strictly excretions of the living substance, that is, of the 
continuous substance. 

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Substance differentiation as seen in Protoplasts was illus- 
trated above by a specific description, for which Amceba proteus 
was used. For differentiation of the living structure as found 
in Metazoan organisms, a starfish egg watched through its 
development is, perhaps, as beautiful a typical illustration as 
the student can have. As it lies in the water before matura- 
tion with its nuclear sac still perfect, it presents to the eye a 
mass of living substance having already area within area of 
pellicular and alveolar organization. Thereafter the history of 
development is truly a record, from moment to moment, of 
more or less unstable permutations, transmutations, and meta- 
morphoses of such structural differences, having complex, 
rhythmical relations to the substance as such, to the substance 
as organism, and to the life history of the race substance. 

At such a time as I have chosen, there is seen within the 
outer protoplasmic pellicle of the egg the mass of general cyto- 
plasmic substance, where changeful rearrangements of the 
foam vesicles arise and pass like dissolving views. The 
vesicles of Butschli's structure have seldom a spherical, but 
rather an irregular contour. They are drawn out of shape still 
more from moment to moment by local tensions in the inter- 
alveolar foam. 

Within the general cytoplasmic area comes the nuclear area 
surrounded by its own pellicle which, in those cases examined, 
was formed of very small vesicles bounded on either side by 
pellicles of continuous substance. It was most like those 
intra-cellular formations which later form the new cell walls. 
The surface of this nuclear membrane yielded at times under 
favorable optical conditions finer vesicular contours which 
produced the " shagreen " effect. 

At this stage, the general nuclear area is a much less dense, 
and optically much more fiuid-seeming, area than the cytoplasm. 
That it is actually so was proven by pressing the nuclear con- 
tents out into the water through a mechanical rupture of this 
membrane and the egg membrane. Yet this nuclear area 
contains local modifications of its vesiculation, which have pro- 
nounced viscosity over that of the cytoplasmic area even. 
These are variably numerous, smaller, spherical areas, which 

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seem to repeat in minute the characters of the whole area, for 
they are surrounded by a pellicle of similar formation to the 
nuclear pellicle, and they contain yet other areas of the same 
kind as themselves, and these even others, so that the whole 
nuclear area is a very complexly dififerentiated foam. 

Within the small spheres appear peculiar optical and physi- 
cal modifications of the interalveolar substance, in the form of 
minute very highly refractive and dense masses. This is the 
" chromatin " substance. It is seen in greatest quantity in 
one, two, or even three of the largest spherical areas, but is 
sometimes found also in very small quantities in several others, 
or in one only, according to stage of maturation. 

Two or more of the largest spheres, which contain the most 
of this substance, approach and coalesce with each other, until 
there is formed one sphere far larger than any of the others, 
and now optically very distinct also, because of the amount of 
the refractive chromatin it carries. 

Some of the smaller areas break up and disappear from time 
to time, and so the whole nuclear area undergoes a thorough 
and continuous series of redistributions of its elements, many 
of these changes being carried on simultaneously. 

The largest sphere becomes the egg nucleus which after- 
wards receives the sperm. Within it also, until the nuclear 
membrane is dissipated, goes on a ceaseless shuffling of the 
elements ; the smaller spherical areas being thus changed into 
a more uniform arrangement which is bounded by the single 
pellicular membrane, and the chromatin substance being dis- 
posed in a seemingly continuous line of interalveolar material 
which forms the nuclear optical network. I believe that the 
rearrangements of the nuclear elements are continuous, and 
that there is never what may be termed a truly resting or 
quiescent stage. The phenomena become at times simply less 
obvious, that is all. 

Even after dissipation of the nuclear membrane, and after 
perfecting of the new nucleus, so far as one can trace the facts, 
there remain small portions of the chromatin scattered about, 
or massed together, in the nuclear substance, and later in the 
cytoplasm. The presence of these was first noticed at Wood's 

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HoII in 1 893, and afforded an explanation to me of the results 
then being attained by other investigators in fertilization of 
eggs from which the new nucleus had apparently been shaken 
entire, and also of supposed non-nucleated fragments of eggs. 
What the final fate of these chromatin fragments was I did not 
determine, being then at work along other lines. They might 
later have been carried to and even into the nucleus, or they 
may have had some special mission of their own in the coming 

It is not possible to describe, for it is not possible to grasp 
mentally, in even their grosser relations, all the changes which 
take place during the developmental phenomena so roughly 
indicated above; for the changes are so continuous, in their 
subtle modulations, and yet so kaleidoscopic in their effect 
on the mind which fails to follow, that directed, or even 
passive, attention is beaten at the outset. And then attempt 
at description is futile, for how can this, with its inescapable 
limitations, be applied to activities which are freely simul- 
taneous in three dimensions of space? To try to grasp all, 
or even most, of those in a limited area is to be utterly baffled. 
Many years of observation could well be given to the changes 
which come about in formation or maturation of the nuclear 
area alone. 

Dissipation of the nuclear pellicle is preceded by contractile 
waves in it, and later the component vesicles, loosing their hold 
upon each other, are carried away and mingled amongst the 
surrounding fluid substance ; for the cytoplasm becomes for a 
time very fluid, from having mingled with it the watery contents 
of the nuclear sac. Later, this local fluidity is much decreased, 
and there is a general gain in viscosity of the whole internal 
portion of the cytoplasm, with marked increase in this respect 
towards the periphery of the egg.^ 

[54] Following up along the same lines the phenomena of 
fertilization and of cell-division, these were seen to be but more 
obvious stages in a long and never-ceasing series of redistribu- 
tions and reorganizations of the foam elements of the living 

1 See Acdvides — filose. 

* See above rhythms of Tiscosity in starfish and sea-urchin eggs. 

Digitized by 



substance of the egg. And it became evident that in the long 
and intricate series of these phenomena, one must learn to 
attach to each area in which is seen true organization, however 
fleeting, of the elements, the value of a true physiological area 
of the mass; and to rank it as a true substance organ.^ 

The very transparent nature of the larval stages of starfish 
affords large opportunity for study of the origin and primitive 
nature of many later substance organs. In such transparent 
forms as the Rotatoria one has an almost limitless source of 
knowledge on these points. Other creatures yield still more. 
From what is found in these, comes illumination for struc- 
tures and processes less easily traced in less accessible proto- 
plasm, which should enable us better to interpret results in 
preserved material. 

[$$\ In all nervous or transmissive areas examined, was 
found an organization strongly akin to that of contractile areas, 
only still more emphasized in a peculiar way. The structural 
characters here belong peculiarly to the finer foam, whether 
the area involves a structure of Biitschli or not. Hence the 
true nervous or transmissive substance forms fibrils of continu- 
ous substance, which may be joined with, or woven into, any 
amount of combination by other modified or unmodified trabe- 
culae of the structure of Biitschli or of a coarser alveolar struc- 
ture. Where these tissues are complex, the appearance given 
by most reagents is singularly false, producing an unnatural 
vacuolation which often simulates a beautiful, distinct structure 
of Biitschli. 

[56] So far as I have seen in transparent aquatic creatures, 
the termination of all nervous or transmissive areas is in the 
continuous or interalveolar substance of a visible structure. 
This does not, as will be shown, preclude possibility of invis- 
ible filamentous extensions into alveolar inclusions, but all the 
direct and indirect evidence is for a special modification of 
the continuous substance simply ceasing at some given point 
in this. 

The continuous substance is sometimes much swollen in 

1 See Activities — Cell-division, in this connection ; also New Formula for the 
Living Substance, and Fosterhood. 

Digitized by 



single or isolated fibrils, tapering down towards its termination 
in the continuous substance to a less and less perceptible 
filament which clearly must end in the continuous element 
of the finer structure of the interalveolar foam. In other 
instances the fibril, or nerve line, of continuous substance is 
uniform in width throughout its entire length. The walls of 
the vesicles in most transmissive areas seem relatively very 
thick with regard to size of inclusions. In some ganglia 
examined the vesicles were, per contra^ large with thin walls, 
and the whole area had a spongy appearance. Transmissive 
inclusions appear to be homogeneously fluid, like those of all 
true ectosarcal formation. 

Between the sacs in which lie the nematocysts of Hydra, were 
seen delicate lines of alveoli which connected these in groups 
of variable number. The alveolar lines were in direct continu- 
ity, as to their continuous substance, with that of the alveolar 
pellicles forming the sacs. The optical aspect of the lines, 
which are to be resolved by the highest powers only, is exactly 
that of such areas as form the skin nerves and other fibrillar 
nerves in rotifers, etc. Watching a Hydra under stimulation, 
those groups of cysts between which I could trace this living 
connection, seemed to coordinate during discharge of nemato- 
cysts from that portion of the animal. 

[57] In such transmissive areas I have seen no direct evidence 
of contraction, but that does not by any means negate the possi- 
bility of such.^ 

[58] Transmissive areas seem to be very stably organized in 
the cases where they appear as special protoplasmic structures. 

[59] It must be noted here that all ectosarcal formations are 
highly transmissive in an organized manner ; differing in the 
matter and manner of transmission according to their specific 
structure. Wherever there is organized contraction there is 
also a certain sensitive transmissiveness which seems to be 
inseparable from such organization of the elements in homo- 
geneous and specially continuous manner, and without breaks 
in the general state of the continuous element. 

[60] In all ectosarcal formations which show marked struc- 
1 See Contractility. 

Digitized by 



tural reduction of the peripheral portion, there appears to be 
increased irritability to stimuli of all sorts, the tactile powers of 
the substance in the widest meaning of the term receiving 

[6i] The optical quality and structure of nerve lines of the con- 
tinuous substance is extremely like that of the "granules," and 
like " chromatin " in all its different groupings as inclusions of 
the finer foam. The similarity of staining reactions of both 
these structures and also of nerve tissues is another likeness 
between them. In actual nerve lines of continuous substance 
I have not seen granules present as such. All these structures 
alike are more or less unstable organizations of the interalveolar 
foam, and can by redistributions along its lines become imper- 
ceptible as distinct areas, though they seem then to impart a 
characteristic optical quality to the whole area in which they lie 

[62] To all of these there must be conceded the value of sub- 
stance organs, as well as of organs of the organism ; and the 
full significance of protoplasmic habits of self-expression. 

As to those granules which, either in their specific form or 
in more finely diffused states, are inseparable comrades of the 
living substance, one is constrained to impute to them some 
direct relation to constant and characteristic activities of the 
substance in which they are imbedded, — that is, the continu- 
ous substance, for of this they are at all times inclusions, and 
at some times compound inclusions. 

I could not determine whether the peculiar optical character 
of these inclusions were due to their included substance, or to 
the character of the continuous substance forming their walls 
and possibly a reticulum. I am inclined to think, however, 
that their refractive quality pertains to their form, and their 
marked tingibility pertains to the nature of their included sub- 
stance. Later, when the facts have been given in relation to 
strial modifications of the continuous substance, this will be 
understood as not a mere guess, but as legitimate inference. 

[63] Protoplasmic differentiations may be summed up as of 

1 This will be more fully discussed in a paper now in preparation for the press, 
dealing with end-organs, consciousness, etc 

Digitized by 



two sorts, which are not necessarily separated in a given area. 

(a) They are based on differences in character of the discon- 
tinuous substances or alveolar inclusions ; or on differences in 
character of the continuous element along optical and physical 
paths of the interalveolar substance. 

(b) As to the first class, form, size, and arrangement, also 
physical differences in kind of inclusions, cause optical differ- 
entiation enabling one to separate areas from each other. 

(c) Leaving out of count for the moment the finer foam struc- 
ture, I find areal differentiation which is dependent upon the 
continuous element for its specific optical character, to be formed 
by a physical arrangement of this along lines dictated by exist- 
ing, but variable, foam structure ; with physical differences in its 
quantity and its appearance, the latter being caused by certain 
physical and physiological changes which have already been 
referred to and will be described more in detail in Striation and 

Physical variations of the foam structure, and the disposition 
of interalveolar stuff with its possibilities of difference in rela- 
tion to any given foam structure ; and these things in potential 
combination with the chemical variations ; surely offer a mag- 
nificent plasticity of conditions and causes for evolution to 
make use of. On this basis arise almost innumerable modifi- 
cations of rhythmic, or intermittent, optical and physical em- 
phasis and their associated changes, in form of alveoli, etc. 

[64] With these possibilities even Biitschli's structure offers 
wide range for area! differentiation ; but when, as frequently 
happens, this is carried down to the finer foam and its reductions,^ 
the opportunities become so multiplied as to sate imagination. 

Nor is it found in fact that the living substance has ignored 
the opportunities. 

[65] In ectosarc is seen the type of substance organization for 
extension and correlation of physiological function. The con- 
nection between the two sets of facts is more readily understood 
when it is seen that in just this type of arrangement of the 
elements the active powers of the substance have full and 
perfect physical opportunity for their exercise and emphasis. 

^ See above ; also Contractility and Activities — filose. 

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The constant correlation of the two sets of facts was 
learned during careful study of contractile and transmissive 
phenomena.^ It is in place here to glance at extension of 
physiological powers on the physical basis afiforded by the 
foam-structure arrangement of protoplasmic elements, and 
peculiarly by an ectosarcal modification of this. 

The myriad potential and achieved differences of physiologi- 
cal areas seem to be susceptible of at least partial analysis on 
this basis. 

[66] In all areas whose office is direct displacement of the pro- 
toplasm — that is, whose function may be termed muscular — an 
increment of power, whether stable or unstable, seems to be 
brought about on a physical basis of a given structure. That 
is, more of the structure of Butschli or of the finer foam may be 
involved, with the result at a given moment of a simple mathe- 
matical increase of the energy, which may be represented by + 
or X . It will be explained under Striation and Contractility 
that the contraction may be complex, and the first contraction 
on the structure of Butschli may be continued by succeeding 
contraction based on the finer foam of the interalveolar material. 
In such case, of course, the same conditions extend themselves 
to the secondary contraction ; but it is without structural 
reduction in the primary structure which may be coarser than 

[67] In all areas where the function is indirect displacement of 
the protoplasm, — that is, in transmissive and sensory areas, — 
I have found always so far a marked reduction of the foam 
structure of the interalveolar substance, proved, it is thought, 
by the optical evidences before advanced for such phenomena. 
Somewhat more is to be said on this head in the section on 
a new formula for the living substance, and much more in my 
forthcoming work on transmissive areas. 

[68] The first sort of physiological extension may be termed 
quantitative extension; the second, qualitative extension of 

Below and behind such grosser differences in the foam struc- 

^ See Striation and Contractility ; also New Formula. 
' See also Selection of Environment. 

Digitized by 



ture of the living substance, lie partially, or wholly, concealed 
optically others whose limits we cannot know and whose nature 
is to be inferred to but a very small extent. 

By themselves, chemical differences constitute a mode of 
areal differentiation of wide range within the limits of Biitschli's 
structure alone; and when the same holds true for the finer 
structure it is evident that what we can resolve of internal vari- 
ations in organization of the living substance is, indeed, but 
a small part of all. That actual physical and' physiological 
differentiation of the living substance far outruns our power to 
resolve local characters is certain. Hence optical differentiation 
is not to be taken as the limit, or sum, of organization, or of the 
physiological machinery of the living substance, but yet may be 
taken as typical of this.^ 

[69] In so-called " low " and " primitive " forms of life, the 
substance organization is seen to be very complex, if here as in 
the Metazoa the sum of all areal differentiations be taken as 
the unit of count ; but it is less stable and more fleeting, — 
often, indeed, to the point of evanescence. Grosser structures 
are openly transmuted, whereas in the adult higher forms there 
is a more stable mask of structure behind which the substance 
carries on its unstable processes. 

[70] Phylogeny is seen, from the standpoint of foam structure, 
to'be a direct progression along these lines in this direction, and 
the types which follow each other in supposed evolutionary 
sequence seem to typify the results in gross of such progression. 

[71] In all forms, from the lowest to the highest examined — 
subtly hidden often, it is true, beneath the stable as well as be- 
neath the openly unstable structures — exist fleeting differentia- 
tions. Nowhere, unless secondarily modified so as to lose its 
vital state, is found a protoplasmic structure so stable and fixed 
as regards the substance as such that it does not permit these ; 
nor does such stability of structure involve or imply renuncia- 
tion of the instability of the substance as such.* Take, for 
instance, such very stably organized areas as the contractile 
cuticle of Epistylis, or the delicate skin of the rotifer, or muscle 

1 See next section, Activities. 

3 See pellicles and alveolar layers, above; and next section, on Activities. 

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bands of this, the crab, or even the tadpole or frog, and within 
the limits of the general stability of the whole area, or sub- 
divisions of it, there may be seen to go on local transmutations 
of finer structure and even local modifications of the structure 
of Biitschli, as well as transpositions of the continuous sub- 
stance. Scheme within scheme of simultaneous differentia- 
tions and physiological activities within a given area is thus 
opened up to us. 

\J2\ In all it is the continuous substance and its organized 
relations to the alveolar inclusions which rank first ; inclusion 
organization ranks second, yet may to large extent control the 
phenomena. The true interaction and relations of these two 
groups of elements are made clearer in later sections.^ 

[73] 1*0 sum up finally: in all the organisms examined, areal 
differentiations of the substance, in the threefold character of 
optical, physical, and physiological difference, express organi- 
zation of the substance as such on a physical basis of the foam 
structure ; and each area so formed, however fleetingly, must 
be understood as a true substance organ, within, or without, its 
possible larger relation of organ of the organism. 

I have tried to iterate the above facts in such a way as to 
express somewhat schematically the aspect in which they 
present themselves to various points of view from which I 
approached the substance as such; and they play into each 
other in such a way as to make this seem a most fruitful plan 
of investigation for the student of living phenomena. 

The few facts given here, like all those from which they were 
selected, are so inseparably correlated with the whole scheme 
of substance phenomena that they cannot be even fairly grasped 
until others, equally representative of other generalized substance 
phenomena, are at once within reach of the mind. Yet it is 
necessary in speaking of them to use to some extent the terms 
which all formulate, bespeaking the forbearance of the reader 
until the end of the argument be reached. 

^ See Contractility, and Selection of £nvir6nment by the Substance. 

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Protoplasmic Activities and Cell Division. 

By the word " activities " I must be understood to mean such 
exhibitions of energy as directly or indirectly cause displace- 
ment of the living substance, — as such, as area, as organ, or 
as organism. 

Useful as it seemed to learn somewhat more of visible proto- 
plasmic structure, there proved to be other facts whose impor- 
tance transcended this. There is no doubt that in protoplasm, 
as in artificial foams, there are structural differentiations which 
imply certain physical stress and controls, resulting from the 
mere physical conditions belonging to Butschli's foam structure. 
This may readily be proven by observation and experiment. 
But besides such phenomena, I find in the living substance 
others which not only cannot be directly grouped with them, 
but which even deny the possibility of such an explanation. 
There exists here another set of controls constantly disturbing 
and transcending, yet inseparably bound up with, arrangements 
and equilibrations urged upon the substance by the given 
physical foam nature. 

[74] In other words : The vital phenomena of protoplasm 
were seen to be not so much manifestations of the visible 
vesicular form of the substance, as upon, or through, this. 

(a) The most important phenomena of the living substance 
are really to great measure independent of the visible foam 
structure. I find them to be referable in all cases to activities 
of the continuous substance of Butschli's structure, and to like 
activities of the continuous substance of the finer foams. 

(b) In most cases they can be seen to pertain wholly to the 
interalveolar stuff, and it is to this as organized, living, substance 
that we must look for the clue to a bewildering labyrinth of 
phenomena and possible interpretations. 

(c) In any visible foam structure, negation of such physical 
controls as have been formulated for protoplasm, is always 
expressed by the continuous substance, or is always to be 
directly referred to it at any given moment. 

[75] Perfect as may be the structure of Butschli, or even a 
finer foam, in any area ; stable as it may seem in any protoplas- 

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mic mass ; there is thus given no true assurance that the living 
substance is at rest. Mass pellicles, alveolar layers, and, within 
these, area upon area of structural diflferentiation, may lie seem- 
ingly unchanged from minute to minute, or from day to day, or 
even throughout the rest of the lifetime of an organism, after 
they are once formed; and still a continuous train of substance 
activities go on, resulting in formation of new structures, both 
within and without the mass. 

The coexistence of a stable and perfect structure of Biitschli 
with a host of metamorphosing activities of the substance as 
such ; this forms one of the strongest reasons I would urge for 
preferring optical research upon living material to more indirect 

[76] The continuous substance of Biitschli's structure seems 
to be very active in all organisms observed. It is indeed most 
commonly in a state of flux, or of active contraction. Some- 
times the two states are found to exist interchangeably in the 
same area which may be stably organized, even for contractile 
purposes, as in the cuticle of Vorticellidae, or the delicate skin 
of rotifers. In pellicles, whether internal or external, excepting 
those which have been wholly altered by deposit of mineral 
substance, or chitin, or the like; no matter how stable and per- 
sistent their seeming under inadequate powers, no matter how 
stable as such the structure of Butschli underlying them; there 
are evidences at one time or another, or at most times, of a 
more or less active flux, or of contractile displacement. Areal, 
or mass, viscosity so great as to produce considerable resistance 
to mechanical tension or pressure, seems no bar to unrest of 
the continuous substance. 

(a) Alveolar layers and many other areal organizations of the 
substance are constantly intermitted, or broken up, by such 
instability of the continuous substance, which is comprehensible 
only through knowledge of its finer foam structure. In the 
finer structure itself is again repeated the same set of phenom- 
ena,^ so that it is plain we cannot stop even here satisfied with 
foam structure as an adequate explanation. 

(b) The motion of the "granules" upon the surface of very 

^ See filose phenomena, cilia, etc., below. 

Digitized by 



Stable areas of Butschli's structure, their transportation from 
place to place in the meshwork, and certain marvellous filose 
displays from the surface of stable pellicles covering stable 
arrangements of Butschli's structure, are thus explained ; also 
those phenomena seen in masses and extensions of protoplasm 
whose whole bulk is less than the thickness of interalveolar 
substance in many an area of Butschli's structure. 

(c) The Brownian-like movements of granules and vesicles in 
fluid, living, protoplasm at times, — notably among the Infu- 
soria during certain states correlated with reproductive phe- 
nomena, when the nuclear bodies are broken up into fragments 
or granules, may doubtless be referred to this cause. 

(d) The formation and the dissipation also of larval areas 
and organs are, as will be shown, due to such activities of the 
interalveolar substance ; whether secondarily or primarily is 
yet to be determined. 

\jy\ The flux and the contractions of continuous substance 
are a cause and a means of displacement of protoplasm, not 
only in bulk internally to the mass and also along lines of the 
meshwork and pellicles ; but outside the mass or area into the 
water or other environmental substance. 

[78] Displacement of this latter sort, which at times involves 
a relatively large proportion of the protoplasm in a mass, may 
go on, yet be invisible except under specially favorable optical 
conditions, and, as will be shown, even under these. The dis- 
placed substance because of its minutely subdivided state may 
still remain invisible ; and may later be returned to the mass in 
the same manner that it issued forth, while the observer is 
unconscious of any phenomenon except, perhaps, of change of 
bulk in the mass, — and for this, is there not always possible, 
and ready, a physical explanation of " osmotic changes " } 

An instance of this may gain a little credence for this rather 
startling statement which will however be amply upheld in 
the course of these remarks. About a large Myxomycete, under 
observation for sequence of protoplasmic movements, the water 
was seen to swarm with granules and a vague suggestion of a 
watery material which was in places somewhat flocculent and was 
at first taken to be excreted substance. Doubtless it would 

Digitized by 



fall under the term of "slime" which is so freely used in 
connection with the Protozoa. The granules showed motion, 
but this was for the moment assumed to be " Brownian," or 
to be caused by currents from the pseudopodial flow of the 

After some moments' absolute rest for the eyes, increased 
illumination, with better adjustment of the draw-tube, and also 
some specially fortunate atmospheric conditions, revealed the 
truth. The creature was surrounded by an evanescent, Gromia- 
like network of protoplasmic spinnings, in the filaments of 
which the granules were held. It was the ever-varying posi- 
tion of the protoplasm of this network carrying the granules, 
which gave with insufficient powers, the effect of dissolving 
watery substance mingled with granules in motion. The net- 
work was in ceaseless flux of form and substance, yet the flow 
and motion of the organism was in no wise altered from its 
characteristic manner and appearance by these peripheral phe- 
nomena. With very little alteration of optical conditions, the 
whole set of external phenomena would disappear like a dream, 
and one would see as usual the seemingly smooth and viscid 
surface of the heavy lobose processes and network. From time 
to time the spinning phenomena were locally intermitted, the 
substance returning wholly into the general pellicle, there to 
be indistinguishably mingled with the peripheral protoplasm, 
and so, after a while, to return to the endosarc.^ 

In the face of facts of this nature, it is difficult indeed to 
hold to any faith in surface tension as an adequate explanation 
of the protoplasmic flow. And these facts are not exceptional; 
they are peculiarly characteristic of the living substance where- 
ever its more secret processes can be traced. 

While Butschli thinks amoeboid flow proper to be physically 
explicable by surface tensions consequent upon the mere physi- 
cal form and the argued chemical nature of the substance, he 
confesses with characteristic candor : " I find myself, despite 
my best efforts, unable to apply the same explanations to the 
finer formations, such as free filose formations, of which Gromia 
furnishes a good example." 

^ See also below, filose activities in starfish ^gg. 

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[79] It is these latter, the free filose phenomena, not amoe- 
boid flow, which I find to be most universal, most characteristic, 
and most fundamental in the living substance. 

Amoeboid flow is but filose flow organized over a sufficient 
area to include to some considerable extent the structure of 
Biitschli. Division between the two has been made on the 
arbitrary ground of difference in relative size and shape of the 
mass involved, and on acceptance of the structure of Biitschli 
as the final structure.^ 

Filose spinnings from pellicles are so frequent that they can- 
not be overlooked. To such cilia owe their origin, and it is 
noteworthy that they arise with equal freedom whether the 
pellicle be external or internal to the mass. Cilia are formed 
by spinning from a pellicle of Biitschli's structure, or from the 
pellicle of the finer froth. In the former case they are apt to 
arise just above the apex of each alveolus of Biitschli's struc- 
ture underlying the pellicle, and there is a quite definite ten- 
dency to formation of alveolar layers beneath ciliated pellicles. 
Because of viscidity of the interalveolar substance in such 
layers, preservations give an appearance which observers have 
described as " microsomes," each supporting a cilium. 

Where cilia arise from the pellicular substance of the finer 
froth, they are very delicate, and seem to clothe the surface of 
the pellicle evenly. In their presence it is not possible, 
because of optical interference, to see the finer foam structure, 
but from their number and evenness of distribution, as well as 
their distance apart, it is hardly rash to say that they bear the 
same relation to the finer alveolation of the pellicle that the 
coarser cilia do to the structure of Biitschli. It sometimes 
happens that the coarser cilia arise at points which correspond 
to union of the continuous substance with the pellicular sub- 
stance. I do not remember a case in which the coarser cilia 
were scattered quite irregularly over a surface. 

Cilia often appear to be formed by splitting up of an undu- 
latory membrane. I believe this to be to great extent an 
optical illusion born of the simultaneous and harmonious 
action of many cilia beating in time, then changing to a rhyth- 

1 See Substance as Organism. 

Digitized by 




mic and wave-like alternation. This sequence of actions is 
common in cilia not only when first formed but after any rest- 
ing period later. It is also possible that they are at first covered 
and bound together by a delicate film of protoplasm so as to 
form a striated undulating membrane, but even then they are 
already formed organs and filose products. 

[80] The length of cilia was found in all cases to exceed 
that given them in published drawings. From the Protozoa to 
the Rotatoria, and from these to starfish and sea-urchin larvae, 
rays, cilia, sense-hairs, and flagellate appendages seem to be 
longer than is figured, except in Messrs. Drysdale and Dal- 
linger's researches upon the monads. This was evident to me 
years ago with no powers higher than a % Crouch objective, 
but then it was supposed to be economy of space which had 
led to suppression of the total length. I am now inclined to 
think that these processes are not seen to their full extension 
by most observers, and there has been evidence in many cases 
that the finest extensions eluded my most carefully arranged 
optical conditions. 

[81] Pellicles spin almost everywhere ; and under very adverse 
physical conditions, as one would think. The interalveolar 
substance seems to be irrepressible. What more unlikely 
physical opportunity for such activity than from the inside of 
alveoli of Biitschli's structure } Yet this I have many times 
seen, even during strongly contracted states of an area, which 
meant great compression of the alveoli. And in coarser alveoli 
of fluid endosarc such extensions are not uncommon. 

[82] The interalveolar stuff spins itself out also along paths 
within itself ; that is, protoplasmic processes, like delicate 
heliozoan rays, are extended through the interalveolar foam, 
between the vesicles, just as such rays are often forced through 
gelatinous material of some considerable density surrounding a 
radiolarian. Indeed, the protoplasmic substance is found acting 
and reacting on and to itself, as if it were a true fluid environ- 
ment. In this fact I find one of the strongest arguments yet 
given for accepting the vesicular nature of the substance. 

[83] Cilia, fibres, and undulatory membranes are formed by 
interalveolar substance in the heart of unbroken networks of 

Digitized by 



Butschli as freely as in the water by peripheral protoplasm. As 
soon as their structure is complete and their activities fully 
organized, they function as if in a watery environment, and the 
surrounding substance gives way before their rhythmic pressure 
as a fluid alone would do. And when these structures are 
from time to time returned by invagination to their native 
environment, the protoplasm again yields fluidly to their beat. 
It is after the manner of a visco-fluid that the living substance 
yields also to pressure of adventitious substances, and to that 
of internal filose activities along lines of the interalveolar foam, 
as well as to pressure of contorted muscle bands.^ The change- 
ful viscosity of the substance gives the necessary physical 
opportunity for such phenomena. 

By such interalveolar activity are explained the seemingly 
independent motions of the "granules," and many transposi- 
tions of the nuclear elements. The motion of granules along, 
and also to and from, pellicular surfaces Butschli found himself 
obliged to interpret by supposing the granules to be governed 
by local surface tensions, and to have in this wise an independ- 
ent motion of their own; especially because these pellicular 
surfaces bounded stable and seemingly motionless alveolar 
layers of his special structure. This inference, although a 
logical one if that structure be accepted as the final physical 
structure of the living substance, can no longer be made in 
view of the facts given here, which establish that such pellicles 
are thick, and often compoundly viscid, foam structures them- 
selves, in whose area can be repeated the chief phenomena 
characterizing the masses they as membranes limit. 

That cell walls have a value in protoplasmic masses of mere 
areal arrangements of the finer foam is shown by countless 
facts, some of which are to be given below. It is certain 
that where cells are in close apposition, their walls become to 
great extent common to both, or all, such cells. Granules and 
even pigment, I have seen pass along the surface of adjacent 
cells, with evident disregard of any separation line between 
them as cells. Again, within the common mass of two fused 
cells in egg cleavage, in that area which is central to the mass 

^ See Striation — formation of aster rays, etc. 

Digitized by 



and which becomes rhythmically more fluid, I have even seen 
yolk transferred from one " cell " to the other. In fine, there 
are strong optical reasons for refusing to hold that the cell wall 
is a true isolator of portions of the substance in a mass, and for 
inclining to believe that the living substance may overstep them 
at times in its more intimate functions which as yet we cannot 
hope to trace fully. 

Perhaps the most striking and significant cases of filose 
activity found were in the developing eggs of starfish and sea- 
urchins. Protoplasmic activities of the pellicular substance were 
seen throughout the normal development of starfish and sea- 
urchin eggs, notably the former. The phenomena were observed 
in two different species of each group, had from two widely 
separated localities in two succeeding years. Many camera 
drawings were obtained both of normal and abnormal eggs.^ 

The filose processes were in the main invisible except with 
the 2.0 mm. immersion with the 8-18 oculars. 

The spinnings contained granules in their usual form, and 
also in their reduction form, when they were no longer visible 
except as a changed optical quality of the substance which 
then appeared quite structureless. The vesiculation of the proc- 
esses was much finer, as a rule, than that of Biitschli's struc- 
ture, and in the normal egg the extensions were wholly of the 
finer, pellicular, foam. In most other respects they were typi- 
cally protoplastic, of a radiolarian type. The filose activities 
seemed to be continuations in kind of those which produce the 
tuft to receive the sperm in the sea urchin ; and in starfish 
accompany extrusion of the polar globules. They arose from 
the general periphery of eggs before cleavage and thereafter 
from the whole periphery of each cell as fast as it formed, 
ceasing only where and when cell surfaces were again fused 
with one another. 

The activities were maintained in normal eggs by pellicular 
material alone, or by this reinforced from underlying, interalve- 
olar, material. That it could be as last stated was manifest in 

^ As I have recently published in the Journal of Morphology a detailed 
account of these phenomena, I will refer the reader to that article for fuller 
accounts of many facts more briefly passed in review here. 

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abnormal egg^ where the excess material came, it was plain, 
from an underlying structure of Biitschli, which might be largely 
involved if the egg's condition were markedly abnormal. 

There was no true intermission, except very locally, at any 
time during development up to the blastula stage when the 
activities ceased externally to the mass for a short time, only 
to burst out once more as cilia formation. This organized 
manifestation terminated them peripherally, so far as I could 
see in following larval stages up to formation of the skeleton 
in three dimensions, or to formation of the proctodaeum. To 
the end of my observations, filose activities persisted internally 
as delicate processes crossing the blastocoele and attaching 
themselves to its walls, to ectodermal processes, or to those of 
mesenchyme cells. For a time after gastrulation, the inner 
ends of the ectoderm cells were elongated, and the optically 
wedge-shaped interspaces so formed were crossed by strands 
and webs of filaments. 

There was no intermission in the course of development 
which could be interpreted as sympathetic with karyokinetic 
rhythms, except a momentary cessation near and at the line 
where a cleavage split was to pass. Following in the path of 
cleavage as fast as blastomeres became split off from each 
other, there sprung up renewed spinnings. These filaments 
extended themselves, with or without ramifications and anas- 
tomosis, but always with markedly less of these phenomena 
than belonged to peripheral spinnings, until they first reached 
filaments, or the pellicle, of the sister cell which was being sep- 
arated. Filaments between cells held, as a rule, a more direct 
course, ramifying less than those of the periphery, and having 
a tendency to structural reduction, seen finally as change of 
optical quality. These traits became more or less emphasized 
in them after their first attachment to a sister cell. With- 
drawing their branches, several filaments would fuse longitu- 
dinally with each other to form bands, or bridges, rather than 
attached rays, of protoplasm between cells. At certain stages 
in cleavage the structurelessness and viscosity of aspect in 
these connections increased, that is, at such time as the cells 
drew together again. 

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The processes sprung up close on actual splitting apart of 
the cells, so that at no time was it usual for the products of a 
cleavage to be physically separate from each other, though with 
slightly less efficient optical conditions one could see no con- 
nection between them. 

[84] Continuity was, in fact, hardly destroyed at any point 
of the substance in cell-division before restored by filose pro- 
jections from both the sister cells. 

The phenomena were more profuse in earlier than in later 
stages. In my most carefully preserved specimens I have ob- 
tained but a much reduced number of fine spin processes, and 
these were somewhat altered. No reagent, not even heat, is 
quick enough, nor can one be applied so quickly as not to give 
the sensitive living substance time to react. Even with great- 
est care such processes are liable to be withdrawn during kill- 
ing operations, and further manipulation for sectioning is very 
destructive to these delicate formations which when living 
have the sensitiveness of end-organs and a rapidity of transfor- 
mation almost equal to that of thought processes. On the 
other hand, the reader is earnestly warned against pseudo-filose 
threads and webs which seem to be caused by chemicals and 
section fixatives, — and it is not impossible that slow killing 
may stimulate cells to abnormal or unaccustomed spinning. A 
student must have large familiarity with such processes, not 
only as to habit and type of formation in normal living eggs, 
but as seen among the Protozoa, especially the Protoplasta, 
before risking predication as to appearances found in preserved 
or sectioned material. 

That portion of sister cells which was at some moment 
internal to both as taken together, was usually last to become 
fused, and here the filaments could be longest watched. 

Not only between sister cells of a recent cleavage did filose 
processes pass. In the four-celled stage, the central cavity was 
filled with a web of these filaments, as was also the space 
between the group and the egg membrane, where they were, 
as a rule, more delicate. In the eight, sixteen, thirty-six celled, 
and intermediate, stages, — for I find the cleavage to be by no 
means regular, as my text-books asserted, but to follow a spiral 

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order, — the whole group of cells continued to be surrounded, 
and bound together across the central cavity, by such processes 
as have been described. Constantly extended, or returning, to 
one or another of the group, it was often impossible to know 
to which cell a process belonged, or with which it would next 
ally itself. 

[85] That such physical connection as is established by the 
filose processes is not physical only, is shown by passage along 
them from one cell to another, and even by a palpable inter- 
change between cells, of both granules and granule-bearing sub- 
stance. Since in certain eggs in the 8~i6-celled stage, in which 
the cells had been induced by continued pressure to separate 
quite widely from each other while continuing their filose activi- 
ties, the order of cleavage and arrangement of cells in the char- 
acteristic spiral was not changed, it seemed clearly proven that 
by the filamentous connections there was maintained true cor- 
relation and interaction of cells, notwithstanding a separation of 
their pellicular surfaces. The fact that such was the case was 
noticed and pointed out to Dr. Whitman long before I discov- 
ered the actual means by which the seemingly inhibitory con- 
ditions were transcended. 

[86] The cleavage pore appeared to be closed by somewhat 
amoeboid activities of the cells about it, and also by peculiar 
activities of the spinning sort. There were at times in the 
optical quality of the filaments those differences which charac- 
terize contracting matter. 

[87] In early stages the blastomeres may at times be sensibly 
reduced by outpouring of filose processes, but except under 
favorable circumstances and very good light, the true cause 
escapes the observer. A film of dust, or moisture, or of grease 
from stage fittings, or even from the touch of the human finger, 
on the surface of the sub-stage condenser, is sufiScient to ob- 
scure all filose processes. 

Eggs on which a continuous series of observations were made 
developed into perfect free-swimming larvae. Increase of heat 
up to a certain point stirred up greater filose activity without 
causing the general development to swerve from the normal. 
Beyond this point, which was not definitely ascertained in 

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degrees, the filose activities seemed to drain ofiF the egg's 
energies and interfere with the course of development. If the 
greater heat were continued, great sheets and webs of proto- 
plasm, nearly all hyaline, were given off; and later, the eggs, 
or cells, would be much distorted by displacement of even the 
yolk-bearing areas. Variable quantities of displaced protoplasm 
extended towards the membrane and adhered to this by means 
of processes along which more matter took its way. Immature 
eggs and eggs overfertilized seemed most prone to these orgies 
of spinning, and in instances quite forgot any cellular destinies 
in exhausting themselves by such fruitless activities. The 
immature eggs were most markedly abnormal in filose wa)rs. 
Most observers have noticed a sort of shapelessness and even 
amoeboid changes of form in such eggs. These protuberances 
are seen with highest powers to be but the base of great webs 
of filose activities. In mature and normal eggs the activities 
do not cause the slightest change of optical seeming, except 
that they emphasize, perhaps, the slightly granular look of the 
surface and may have been up to this time mistaken for such 
a condition.^ 

Even more remarkable from some points of view than the 
activities of the cytoplasm just noted, were those of the polar 

[88] From the moment of their extrusion the polar globules 
showed filose activities of their cytoplasmic substance, no less 
marked than those of the egg cytoplasm ; and usually in more 
or less intimate connection with these latter. They retained 
these activities until time of closure of the cleavage pore when 
they were drawn, or migrated, inside the blastula, and there 
retained their social union with the ectoderm filaments until 
formation of the endoderm processes and, still later, of the 
mesenchyme web, when they could sometimes still be detected 
in union with this also. 

Their chromatic substance preserved all the while a most 
sharply marked and characteristic optical appearance, though 

1 The fact that even in these strongly emphasized abnonnal cases the spinning 
activities have not been described, points to the reasonableness of believing that 
the finer, less obvious filaments of the nonnal eggs have been overiooked also. 

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it was often separated in the form of granules and then again 
drawn together into a compact and nuclear-like mass. 

In all respects these strange little masses of living substance 
looked and behaved like small protoplasts. Their webs and 
processes were ceaselessly extending themselves on all sides 
and becoming mingled and entangled with those of the cyto- 
plasmic webs. The intercellular space above which they lay 
and which came later to be the cleavage pore was crossed and 
often well filled with fused, and interlaced and intermingled, 
spinnings from the polar globules and adjacent cells. 

That the polar globules persist for a long time as active and 
seemingly independent substance, or units, was clearly shown. 
More there may be of even greater significance, seeing that 
they are again returned to the general mass of egg substance. 
Is it not possible that, as the cells have but a semblance of 
separation, these bodies also are but portions of the organism 
temporarily segregated for physiological purposes ? 

Protoplasm squeezed from eggs in the earliest stages spun 
more or less vigorously, according to the time in certain 
rhythms of viscosity of the cells that the experiment was 
made. Later, the filose phenomena were replaced by amoeboid 
or lobose processes, or ectosarcal areas, all of which varied in 
their character according to the time in these same rhythms 
and ^n certain other rhythms which pertained to the whole 
development. With regard to viscosity of the substance in 
developing starfish eggs a number of very interesting rhythms 
were broadly noted. 

In the single-celled stage, from the moment the egg is laid, 
there is a steady, progressive increase in peripheral viscosity of 
the mass, which gradually extends further and further inwards 
after the nuclear sac membrane is dissipated and its fluid con- 
tents distributed through the general cytoplasm. 

After this, a quadruple set of rhythms of viscosity were 
noted in starfish and sea-urchin, also in rotifer, eggs. One set 
of rhythms held relation to cleavage of each cell, another to 
the whole progressive development of the mass. In addition 
to these was a double set of rhythms of varying viscosity in 
areas of the mass. One set held relation to position in individ- 

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ual cell-masses; the other to position, with relation to the whole 
mass of the embryo. 

According to the point of time in each of these sets of 
varying rhythms, did the viscosity, elasticity, and resistance to 
pressure, of the mass or area vary ; also the specific reaction 
of the substance when brought artificially in contact with 

From the first it is true, that those portions which lie inter- 
nally to the embryo as a mass are more fluid; and conversely, 
that the more peripheral portions are more viscous and resistant. 

Internally to each cell also, whenever it stands separate, 
except for the filaments, from the mass, or its sister cell ; there 
is always an area of more fluid substance. 

After each cell again becomes firmly welded to the mass, or 
to its sister cell, that portion which is internal to the whole 
mass as then formed, becomes more fluid, without respect to the 
number of cells. 

This seeming sensitiveness of the substance to its position 
in the common mass is the more remarkable since the centre of 
the group is a cavity full of the same watery environment as sur- 
rounds the periphery^ because the conditions then would seem 
to be identical for all the peripheral substance, yet it reacts 
diflFerently at different points with little or no regard to this. 

In the four-celled stage, when the cells are sometimes firmly 
welded together for long minutes, there was marked fluidity of 
those portions, including the pellicular wall, which projected 
inwards towards the cleavage cavity. From this portion the 
filose processes extended, filling the cavity with a protoplasmic 
web. Yet the firmly knit and relatively very viscous peripheral 
portions were no less active in this respect. When a more fluid 
area comes in the rhythm to lie between two sister cells, the 
cell walls also share in the relaxed physical state. 

[89] The viscosity of the substance is, then, rhythmically 
modified according to its temporary position with respect to 
external environment of the whole embryo ; greater or less 
fluidity being assumed in the same region of a cell or the mass, 
according to its position as internal or external to that organ- 
ization of the living substance. (See Substance as Organism— 

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Raphidiophrys elegans,) In the two sets of position rhythms 
it is plain to be seen that, while the substance controls its local 
state and structure with reference to external environment, 
this latter cannot be held to be the controlling factor, but only 
an influencing factor. Besides the above major rhythms, every 
smallest area of the cytoplasmic and nuclear substance seems to 
have rhythms of its own of varying viscosity; but the tangled 
maze of these it is now beyond our power to thread. They bear 
a constant relation to the phenomena of cell-division and karyo- 
kinesis, and are visible chiefly as local or strial modifications. 

The production by protoplasm of a homogeneous, or ectosarc- 
like area, when it is artificially brought into contact with the 
water, was found to vary greatly, both as to amount and kind 
of such a formation, according to the time in the above complex 
set of rhythms in the area experimented on ; and to vary also 
in the peripheral substance, as such, from the internal substance, 
as such. 

With regard to experiments made upon living substance 
crushed from protoplasts, in attempting to confirm, or disprove, 
experiments upon purely physical foams, it must be borne in 
mind, that, in the former case, the substance remains for a 
long time living, and in death disintegrates, so that we can 
not too readily assume a rapid formation by it of ectosarcal 
areas to be due to the same cause as the gradual formation of 
a hyaline peripheral area by the non-living foam ; basing such 
conclusion on an optical similarity between the two formations, 
which is in the main superficial. 

That the formation of ectosarc by the living substance is 
not a direct physical result of contact with water, is abundantly 
proved by the fact that, in the same protoplast, from moment 
to moment, contact with water produces or does not produce 
ectosarc, excepting always a pellicle. Further there are in 
many protoplasts and higher Protozoa also, areas of most 
fluid substance, which on contact with water never form any 
ectosarc further than a very thin pellicular film. In Amceba 
proteus and Amasba radiosa^ the ectosarc may be at one time 
only a film, and then again extend half across the body of the 
animal. It is frequently, also, formed within already existing 

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areas of ectosarc, the rush of substance transforming itself 
here as rapidly as when close to the water. 

That ectosarc is not altogether a mere physical redistribution 
of the elements, but has reference to physical organization of 
these for physiological function, is manifest when one studies 
a transformation of ordinary, irregularly structured, endosarc 
into beautifully organized and uniform ectosarcal areas, in which 
are then seen organized contractile activities. Such phenomena 
can be followed from moment to moment in Amceba radiosa^ 
where the endosarc can be seen to transform itself in a short 
time into ectosarc of mixed structure ; this to pass into a mani- 
festly organized state, and then into contractile states, in which 
all structure may or may not disappear from sight, the substance 
becoming so dense and refractive as to fairly glitter, and so 
resistant that when it has the form of long lash-like extensions 
they may be bent elastically by pressure as if they were bristles, 
and then in another moment will lash themselves about like the 
flagella of //i?/^w»i//^ or flow like the pseudopodia ol A.proteus, 
In such viscous states in A. radiosa when a posterior mass has 
a thick ectosarcal covering, I have seen this thrown by contrac- 
tion into folds which were so refractive as to simulate quite 
well spiculae of flint, and by these contractions the whole mass, 
including an anterior, film-like, ectosarcal flow, seemed to be 
urged forward. In another moment the creature was a large 
fan-shaped film of protoplasm, so delicate as to be scarcely 
visible, preceding a protean, partially naked, lump of somewhat 
filose endosarc. 

In addition to the facts cited and others of kindred import, 
negation to any explanation of cleavage by surface tension was 
had early in the summer of 1893 by the following experiment. 
A starfish egg in the two-celled stage was gradually compressed 
until the pellicular wall was broken and the peripheral proto- 
plasm exuded at points from both cells, forming very short 
blunt processes. Little by little these were augmented by 
outflow of hyaline interalveolar substance, and at the same time 
they showed local changes in structure, and amoeboid activities, 
the substance flowing here and there and spreading itself out 
over the surface of the whole egg. There seemed to be a 

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ceaseless series of these activities going on for hours, the sub- 
stance heaping itself up here and there in lumps, and then again 
forming papillose processes over wide areas, and again, a thick, 
smooth pellicle. 

Little surface tension for the mass would seem to persist 
under such conditions. As a matter of fact, the peripheral 
pellicle itself relaxed in places, extending its substance no 
longer in filose processes but in amoeboid lumps, or knots, or 
films. Yet, all the while, the phenomena of karyokinesis and 
of actual cell cleavage went on ; more slowly than in normal 
eggs, but still without pause. First into four, then into eight, 
then into sixteen cells did the mass divide, forming internally 
perfect walls for all the cells, though there never occurred any 
actual separation of these from each other. Still, in the usual 
acceptation of the term, the cells divided ; that is, their mass 
was subdivided by, so-called, cell walls. 

In the ectosarc formed by the egg substance after being 
squeezed into the water, structural reduction, and then in some 
cases, a reappearance of Biitschli's structure, were observed. 
The finest vesicles which could be detected showed no change 
of contour, so that here, as in protoplasmic ectosarc, a structure- 
less appearance could hardly be attributed to mere stretching 
of the alveolar lamellae, especially as the meshwork became 
often thicker and optically more conspicuous, rather than more 
tenuous and less obvious as reduction went on. 

When pressure was applied to eggs in 4-16-celled stages, in 
which the cells last divided were still optically separated, fila- 
ments connecting the cells were seen to become thicker, denser, 
more refractive, and yellowish in tone. At the same time, if 
the pressure were slow, the viscosity and resistance of the cell 
mass also increased; and the cells showed a similar change in 
quality of their pellicular stuff, while many peripheral processes 
were withdrawn. 

Rapid, or sudden, pressure effected a separation of the cells 
more readily, but seemed to be followed by the same optical 
changes in the cell protoplasm. If actually separated, but 
without rupturing the membrane perceptibly, as was done a 
number of times by pressure of a mixed rolling and squeezing 

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nature, the cells passed soon after through a great change of 
viscosity, visibly relaxing. They then showed rather marked 
change of contour, and afterward renewed their spinnings until 
once more connection was reestablished amongst themselves, 
when by degrees they drew more and more closely together 
until they touched. The walls then coalesced and the two, 
four, six, eight, twelve, or more, cells were again a solid mass. 
After such coalescence there came on always a marked increase 
of viscosity in the whole mass, and markedly greater resistance 
to pressure. Just before renewed division this would again be 
somewhat relaxed, and the spinning phenomena more active. 

During the past two years I have observed in the blastomeres 
of frogs' eggs the same increase of resistance under gradual 
pressure, and similar changes in some few of the lobose bands 
of protoplasm or ectosarc by which, when separated, they had 
restored connection. 

[90] There can hardly be a doubt but that there is here shown 
a definite physiological resistance to certain adverse mechanical 
conditions in environment ; that the living substance responds 
in character of its own powers to stimulus of a given sort ; 
that this response is to conditions which are probably new to 
the substance, and is, moreover, contrary in its nature to that 
given by purely physical foams. 

For, had the substance thus dealt with been an inert, purely 
physical, foam, bridges and bands of it connecting the cells 
would have shown a most contrary series of reactions. Under 
pressure that forced apart the cells, they would have become 
more tenuous and longer, as well as less distinct and less 
refractive optically, and the pellicular substance also would have 
become stretched and less refractive. The peripheral processes 
would have been increased in length and mass, and also, no 
doubt, in number. 

The phenomena of the living eggs were on the other hand 
exactly such as characterize all protoplasm, whatever its form 
or position, in contractile states and especially during active 
contraction. Further proof that this was actually the case was 
given during the usual drawing together of separated sister 
cells after cleavage was ended. First the cells usually rounded 

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themselves up on all sides, except those of the cleavage plane 
which were connected by filaments. Finally, after a number 
of changes of form, these sides, which were then rather flat- 
tened, approached each other more and more, while the 
protoplasmic bridges showed greater homogeneity and even 
struct urelessness, with increase of viscosity and refractiveness. 
They thickened too, in many instances, as the cell surfaces 
drew together, until at last the pellicular surfaces were too close 
to admit of further observation. 

In sea-urchin eggs, the spinning phenomena were fundamen- 
tally the same ; the filose processes, however, were less profuse 
and finer. Similar varying rhythms of viscosity, and flux of pel- 
licular and interalveolar substance were also observed. In this 
form the segregation at certain rhythmic intervals, or rather in 
certain rhythmic progressions, of pigment granules from certain 
portions of the cytoplasm was interesting and suggestive, not 
only in connection with a flux of continuous substance along 
lines of the meshwork and pellicles, but as correlated with 
structural preparation of certain areas for specific physiological 
function. Before each early cleavage, pigment granules were 
carried along the pellicles in a flux of substance towards the 
line where the split was to take place, and appeared to be 
carried onwards and inwards from this point, but about this last 
there is yet some doubt in my mind. 

In rotifers' eggs a similar segfregation of yolk is extremely 
marked, the very planes of cleavage being thus for a long time 
clearly indicated before there is any other sign of an order of 
cleavage among the cells. 

Such phenomena are thoroughly in accord with the general 
habit of the substance in withdrawing interalveolar substance 
to an area where its activities when organized will not be ham- 
pered by inelastic, or irregular, inclusions ; or of withdrawing 
the inclusions to another place. The areas cleared of yolk in 
segmenting eggs are noticeably areas in which are about to 
take place organized contractile activities, as amphiastral areas, 
or cleavage planes. 

Pigment, yolk, and granules are carried thus in instances 
from cell to cell. Flux of interalveolar and pellicular substance 

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can be traced often by movements of such inclusions. That 
cell walls form at times no barrier to wanderings of the living 
substance seems to be shown. It is noteworthy that these things 
can take place without disturbance of a structure of Blitschli. 

Since these observations were made, there have been many 
additions to our knowledge of intercellular connections in 
higher Metazoa. They are but links in an endless chain of 
evidences that the living substance clings pertinaciously to its 
protoplastic habit; and that the formation of a firm peripheral 
pellicle about subdivisions of a whole mass is in any case no 
proof of actual physical, or physiological, separation of the 
substance, any more than the presence of stable structures of 
Biitschli argues a quiescent state of the living substance. 

I am aware that, because of the restrictions of the preserved 
and sectioned condition in which most intercellular connections 
have been discovered, these are commonly regarded as stable 
processes from the cells. I think we have now grounds for 
thinking of them as possible routes of travel of. the unstable 
and migrant substance, or at least as protoplastic formations. 
For everywhere I find the living substance shifting its where- 
abouts, combining and recombining its elements, and always, 
after each turn of the physical kaleidoscope, there is found 
associated with the new design a new manifestation of ordered 

Clearly it is a habit of the substance to be tenacious of con- 
tinuity between portions of its mass, except when it organizes 
its activities and itself to transplant a portion of itself to a new 
environment.^ Those apparent contradictions of this fact 
offered by blood corpuscles and wandering cells do not interfere 
with this view since in the case of the latter, and indeed all 
cases in which exist protoplastic activities, there is reestablished 
from time to time by means of these a true connection with 
the areas of viscid foam through which they wander ; and in 
case of the former, a slight readjustment of our standpoint, or 
even a slight addition to our knowledge of their true life history 
along the lines of these studies, may bring them strictly within 
the same assumption. 

^ Fosterhood. 

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[91] The true nature and mode of formation of cell walls seem 
to be that they are but pellicular modifications of the continuous 
substance of masses into plates, or membranes, which are as 
readily formed internally as externally ; that they do not differ 
either in origin or constitution from other internal and external 
physiological pellicles or ectosarcal formations ; that they are, 
in short, not substance dividers nor substance isolators, but 
substance strictures, substance differentiators, anddifferentiatiom, 
substance organs; and finally, that they belong primarily to the 
mass and but secondarily to cells, 

[92] We can no longer regard these formations as having for 
the living substance the value of a prison wall. We must rather 
look upon them as substance strengtheners ; as devices for 
securing a qualified independence for areas which yet maintain 
absolute physical and physiological continuity. They limit to 
some extent the areas surrounding centres of control distributed 
through the mass. By such "centres" I mean the nuclei, which 
would seem to control the supply and the use of specific 
environment of the substance as such. 

The pellicular membrane which surrounds cells is no better 
reason for thinking of these cell areas as isolated, independent, 
reproductions of the original unit, than is the nuclear membrane 
for calling that organ an intra-plasmic cell. 

[93] Notwithstanding this reasoning, Metazoan develop- 
ment may still remain to our thought a multiplication of cells, 
but must cease to be to it, as of old, that multiplication of 
morphological units which Huxley preached. It is rather sub- 
division of the substance, with or without growth of mass, than 
a reproduction of the first egg cell. Cell division is mass differ- 
entiation ; that is its true meaning, — making it one with all 
the host of substance organizations of the elements of the living 
substance for physiological function. 

Passing in review in the light of all my facts those colonial 
and compound organizations of living substance found in both 
Protozoa and Metazoa, it becomes evident that so long as there 
is between the units composing it, a protoplasmic connection, 
even the most delicate, a whole colony must be conceded the 
value of a morphological unit, and each so-called cell, or unit, of 

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the colony must be looked upon as merely an areal organization 
of the substance of the whole. For there is no true difference 
between a connection such as is shown in Vorticellidae, where 
the contractile stalk of each unit is continuous with all the 
others by way of a common or multiple stem ; and that seen 
to be established in a compound Heliozoan, or compound 
Ccelenterate, or even in a developing starfish egg. 

It seems strange that Huxley when he had been led to 
declare for the compound Coelenterates the value of true mor- 
phological units should have missed the unity of such a relation 
throughout the living kingdom.^ . 

. [94] To sum up very broadly what has been shown of the activ- 
ities of the living substance : these have been seen in all cases 
observed to be correlated with an ectosarcal arrangement or 
organization of the two groups of protoplasmic elements ; and 
they have been seen in all cases to transcend any visible struc- 
ture. In other words, the activities of protoplasm are not to 
be directly referred to, or accounted for by, any structure we 
can resolve by our present optical resources. 

[95] Further, all visible structures have been seen to be a 
direct result of activities having the same character as those seen 
to be inseparably correlated with the visible foam structure ; that 
is, visible structures are the direct result of activities correlated 
with a finer and not visible series of kindred structures, in 
which we are constrained by the evidence to believe the same 
relations of living substance and structure hold good. Thus 
any explanation of the activities by structure is removed beyond 
our present reach. 

[96] Last and most important, the visible structure of organ- 
isms, of organized masses of the living substance, was seen to be 
relative only ; that is, the areas exist in relation to each other 
and to the organism as such, while the living substance in g^eat 
part ignores these visible fixities, circulating freely beneath the 
mask they form for us. In its more secret life protoplasm is 
unstable and protoplastic, however rigid the structural forms it 
maintains. The form persists — the substance composing it is 
not fixed at any point in it. 

^ See Substance as Such and as Organism. 

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True protoplasmic life and habit lie hidden beneath those 
very forms and habits of a grosser nature in which they have 
hitherto been sought ; as too often happens, the oracle has 
been veiled by the very terms of its expression. 

In those sections which follow after Contractility, many more 
facts are added and this line of thought is further expanded by 
their means. 


At the outset of this record of facts, I should like to disclaim 
any attempt to deal as one who knows their conditions and 
limitations, with the great problems of molecular physics in- 
volved. Biitschli has proved upon his artificial foams certain 
physical forms and interactions, and applied these in his re- 
searches into the phenomena of the living substance. I have 
ventured no more than to extend still further along the same 
lines the same set of comparisons, using however certain other 
phenomena which escaped Biitschli's observation, and seem not 
to have been recorded at that time for his criticism. 

In supposing that I might follow so far as he essays to take 
his readers, I may have been misled by the very lucidness and 
thoroughness of Biitschli's analysis and demonstrations, so far 
as these go ; and for precise understanding of them I have 
depended to some extent upon the authorised English transla- 
tion by Minchin. 

Biitschli found that a linear arrangement of alveoli, in both 
fluid and very viscid foams, produced an optical, or what might 
rather be called a psychological, striation of the substance. 
He has shown that this effect is given also where the alveoli, in 
addition to this purely mechanical arrangement, are elongated 
in the same direction by compression, or osmotic changes, or 
by tension of any sort causing extension, or stretching, of the 
lamellar material. He finds himself able to group under these 
heads all optical striation of the living substance, and he ex- 
presses himself to the effect that a fibrous appearance depends 
solely upon the arrangement, or extension, or stretching, of the 
meshes in a given direction. Irregular, and tangled, fibrous 
structure, can always, he thinks, be explained by the fact that 

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at these points simultaneous or successive strains took effect 
in different directions. 

Here as elsewhere I find that Biitschli has laid down an 
impregnable basis of fact. I find optical striation to be, as he 
asserts, associated always with a distinctly linear arrangement 
of the meshwork substance of a vesicular structure. I find 
striations like those he describes as typical in his artificial 
.foams, are plentiful and can readily be found in living proto- 
plasm. It is easy to produce them also in almost any area 
of the living substance by simple compression, or extension, 
between cover and slide. The best results are to be had in 
areas quite but not too viscid. 

I find, beyond this, that there are many conjunctions of cir- 
cumstance which constrain or impel the mind to strial interpre- 
tation of an optical network. Some of these seem to be very 
slight causes, one would think, but the ever too ready mind 
receives impulse from them, nevertheless, in one direction rather 
than another, and, moved by such slight matters as relative 
size, or even difference in geometrical shape, of vesicular con- 
tours, hastens here or there, shaping lines of emphasis for itself. 
The accustomed haste of its movements, being now under the 
spur of enforced attention also, strengthens such an impression, 
and may conduct the eye in tortuous or broken curves, as well 
as in more or less straight lines. In some areas, influences such 
as these urging the mind simultaneously along a number of 
paths in a given plane, contend with each other, and so pro- 
duce an aspect of tangled fibrous structure. 

To hold well in leash, without unduly restraining, this eager 
poise of the mind ; to protect it as much as possible from the 
vitiating habit of predication which ordinary education and 
human intercourse teach; these are some of the most necessary 
and most fruitful lines of self training for those who seek to 
know the living substance as such. Where the mind follows 
within the limited depth of focus of the high powers used, it 
selects from amongst the fused lamellae of the very irregular 
polygons which at times represent the foam structure, the most 
obvious and connected paths or lines of the continuous sub- 
stance ; and a falling of these above or below the plane of an 

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optical network,- makes such false st nations which really have 
little to do with the true and common structural relations of a 
strial sort. In sections, a tearing, or displacement in shrinking, 
of the visco-fluid lamellar substance may cause a marked, but 
false, seeming of structural striation. However strongly mesh- 
work lines extend in a given direction, caution must be used in 
inferring tension, by osmosis or what not, in that direction, for 
a marked, and seemingly simple and direct, striation in one 
optical plane may express diverse strains in other directions. 

In tracing many optical striations of protoplasm along lines 
of, so-called, optical emphasis, I presently became aware that 
besides these were others of a different origin and nature, 
having marked specific characters. Here too it seemed that 
though a true word had been spoken and a strong word, yet, 
as in the case of the foam structure, it was not the " master- 
word " of the situation. Comparing the new sort of striations 
with those just described, it seemed to me that certain defi- 
nite physical, and psychological, or optical, qualifications of 
these latter did not apply to the former. If I am mistaken in 
supposing these not capable of being grouped with products of 
purely physical, or mechanical, lamellar extension, the physi- 
cists will not leave the error long uncorrected. 

Such striations have a basis of actual physical difference in 
mass and character of the continuous, or interalveolar, sub- 
stance, which causes them to have marked optical emphasis. 
Many of them were correlated with contractile activities, and 
were found to depend not only upon a characteristic passive 
arrangement of the continuous substance, but upon intermittent, 
or rhythmic, states and physical modifications of this. These 
were manifested optically by rhythmical, or intermittent, changes 
m its mass, density, refractive quality, viscosity, and position 
in the mass, or area. Striation may exist before and after con- 
tractile function, but the latter always produces in the substance 
either optical striation of a visible structure ; or optical changes 
which are to be correlated with a similar structure beyond our 
reach ; or increased emphasis of an existing striation. 

There is another striation which may exist without being 
associated with organized contractile function ; this is a local 

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difference in mass, or quality, of the continuous substance. It 
is often due to its interalveolar filose activities, but may result 
from more stable local increment where the interalveolar foam 
has a secretory or irritable function. 

Contractile striation, and even true fibrillation, appear some- 
times in very fluid areas of protoplasm. Then alveoli will be 
drawn into line, for the moment at least, along the course of 
the optical striation. Many swift and evanescent contractions 
of the fluid substance in protean forms or areas can thus be 
traced, and causes and means of motion of the protoplasm of 
amoeba and kindred forms can best be seen and followed. 

The line established by a contraction wave may be very tor- 
tuous among the vesicles, and thus bring into line, by, or for, 
organized action, those which were at first quite far apart. The 
converse is obvious, and the value of such a method in forming 
compound areas having enormous extension powers in given 
directions must be great. 

It has already been stated that, from time to time, in areas 
and masses, the living substance varies its viscosity, and hence 
its optical qualities, becoming denser and more refractive 
when viewed with powers too low to resolve structure. In 
examining such protoplasm wherever possible under powers 
adequate to resolve all, or part, of the foam structure, the physi- 
cal modification was found to be due to like qualities in the 
continuous, and chiefly in the interalveolar, material of the 
visible structure, irrespective of size, or character of arrange- 
ment, of the alveolar inclusions of Biitschli's structure. And 
the continuous substance held these characters for just so long 
as the area, as a whole or locally, held the optical characters 
noted. And wherever, under high powers, the interalveolar 
substance showed such characters in any area, however small, 
that area under lower powers had a marked distinctness and 
refractive quality which were persistent, or intermittent, in 
sympathy with the same character in the continuous element. 

Further, those parts of the continuous element which had 
more marked refractive quality, were visible as distinct structures 
under powers which showed nothing of the rest of the network, 
however one might isolate minute portions for examination. 

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Conversely, wherever there were seen such refractive portions 
of lines of continuous substance, I either found directly by 
change to higher powers, or, where the limit of magnification 
had been already reached, was able to argue from other activi- 
ties of the same area, or from associated optical phenomena, a 
true foam structure. 

It frequently happens that there is an alveolar arrangement 
linear in two directions, and that parallel lines of an optical 
reticulum cross each other at sharply defined right angles, or 
at equal angles of other sorts, yet in these cases there may be 
true and very marked strial emphasis upon one set only of these 
lines ; or may be upon both in alternation ; or equally upon 
both, being in this last case markedly different from surround- 
ing optical network lines also caused by linearly placed alveoli. 

In both Protozoan and Metazoan groups, striations associated 
with contractile activities appear singly, and in simple or com- 
pound groups. Even so comparatively simple a form as Vorti- 
cella has a footstalk in which I find no less than five different 
areas of organized foam structure grouped in physiological 
alliance, each bearing some special part in the contractile ac- 
tivity ; and there are indications of even more such differentia- 
tions beyond our reach. 

It is often impossible to resolve with striations, cross lines 
of an alveolar network, simply because the striae have been 
made visible by increase of mass of the interalveolar substance, 
— with increase also perhaps of its refractive power along those 
lines, — in a vesicular structure whose lamellar substance, with- 
out such increment, were far beyond our reach. Sometimes 
also, striations coincide, not with an optical reticulum of the 
plane in observation, but with that of a plane at right angles to 
this. Such a case was found in the cuticle of Stentor cceruletts. 

If strial or fibrillar appearances of contractile areas, as cor- 
related with physiological function of the substance there, are 
grasped in relation to Butschli's structure, it becomes pos- 
sible to understand how these results apply to phenomena 
found in the finer structure. 

As a typical instance of contraction organized on a basis of 
Butschli's alveolar structure, I will take the cuticle of Epistylis 

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flavicansy which has been variously described from time to time. 
Earlier observers thought the cuticle to be decorated with a 
beaded formation in the peripheral protoplasm. Greeff later 
ascribed the optical appearance to two sets of muscular fibrils 
surrounding the creature and crossing each other more or 
less at right angles. He thought that these two sets of fibrils 
contracted alternately as the animal extended, or shortened, 

I find the true structure to be that of a large cuticular layer 
of vesicles of Biitschli's structure, quite uniform in size, and 
covered by a pellicle of protoplasm which in its turn shows a 
peripheral layer of delicate alveoli of the finer foam, (now for 
the first time described). The interalveolar substance of the 
cuticular layer is much modified from mere lamellar substance. 
It contains strands, or plates, of the finer foam, which pass 
rhythmically from viscid to fluid states, showing from minute 
to minute a host of local, intermediate states and modifications 
of its structure. Such modification of the interalveolar sub- 
stance is along lines in one plane, and in two directions. The 
lines lying between the alveoli intersect each other at such 
angles as the shape of the body directs. 

When the animal was in medium extension, the optical stria- 
tion was about as marked in one set of lines as the other, in 
neither so marked as during more extreme change of shape. 
The effect was then somewhat strongly that of the muscular 
fibrils Greeff described. If greater extension occurred, that 
set of lines parallel with the increasing diameter grew thicker, 
and gained in optical emphasis. At the same time, the other 
set of strial lines, running parallel to the decreasing diameter, 
lost optical distinctness by becoming thinner and thinner as 
extension proceeded. When the animal contracted, the phe- 
nomena were reversed. But still the loss of emphasis was 
always parallel with the decreasing diameter; and gain in 
optical and physical definiteness was always parallel with the 
increasing diameter. 

If Greeff s idea were the true one, and the strise represented 
two sets of muscular fibrils, quite another set of appearances 
would be seen, for then those strise corresponding with the 

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shortening axis of the body would thicken as they shortened ; 
while those having- a transverse direction would of course thin 
as they elongated with the mass. The alveoli too, would then 
become more widely separated as the fibrils shortened, and 
woidd draw nearer together as these elongated. But it is just 
the converse of this that I found to take place in Epistylis, — 
for there the alveoli approach each other along the shorten- 
ing, and become more widely separated along the lengthening, 

If we attempt to explain the phenomena simply by change 
of shape of the alveoli under tension stress, as in Biitschli's 
tentative hypothesis of contraction, the conditions exacted are 
just those needed in Greeflf's explanation, and hence cannot be 
in place here. Especially are these things true, if the pres- 
ence of the finer foam be taken into consideration, as indeed 
it must, for, during relaxed moments, the interalveolar sub- 
stance showed filose phenomena, extending itself into the 
inclusions of Biitschli; the alveoli regained locally more 
rounded contours during general contracted states ; and the 
strial substance even split into strands, or became visibly, 
though very minutely vacuolate ; and during local contractions 
in these strands the alveoli slipped slightly along the striae, 
or had their optical network drawn into triangles, or other 

Closer observation showed that the increased emphasis in 
the striae was due to an actual displacement of interalveolar 
substance during contraction. During displacement of the 
mass in any direction, the interalveolar material forming the 
last emphasized set of striae was in great part displaced, 
the displacement being an actual one, and following, as it 
seemed, lines of least resistance between the yielding alveoli 
filled with fluid ; which may be prepared paths. The contrac- 
tion wave which urged forward the substance, drew together 
the alveoli, and as the activity had the usual wave-like prog- 
ress, the cross striae became progressively increased in mass by 
access of interalveolar stuff. The ectosarcal mode of organi- 
zation of the elements as typically seen in this cuticle of 
Epistylis, and which is characteristic of all areas of organized 

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contracting substance, is now seen to have manifest physical 
advantages for such a set of physical changes, controlled by 

These phenomena were found gradually to be a key to a 
whole host of contractile phenomena of the living substance, 
as seen in optical changes of a visible, or inferred, vesicular 
structure. All apparent contradictions were found to fall 
under the same explanation, when based on the finer struc- 
ture of the interalveolar substance itself; wherever the con- 
tinuous substance in contracting areas was seen to thin as it 
elongated in sympathy with mass extension, and to thicken 
in sympathy with contraction, it was found to be itself the 
organized foam, or area of contraction. That is, the sur- 
rounding substance bore but a passive part in the contraction 
phenomena, and the striae,. if there were more than one, acted 
in harmony but without that structural dependence seen in 
the Epistylis cuticle. In such cases, then, the structure of 
Blitschli, or the visible organized structure, was not the active 
basis of physiological function. 

In the highly elastic cuticle of certain rotifers, Philodinaea, 
where there is a beautiful organization of the elements as 
in Epistylis, it was noticed that striae, after becoming, as in 
Epistylis, thicker and more obvious until the alternate set 
of lines was almost obliterated, then began to grow thinner 
and still more highly refractive, while' extension of the mass 
and of their length continued. In other words, the limit of 
displacement of the interalveolar substance upon a basis of 
Butschli's structure being reached, extension was prolonged 
upon a basis of the finer foam; the strial substance then 
functioning as true fibril. 

All such cases, whether stable or intermittent, or only 
occasional, fall under the head of muscular fibrils of the 
substance. There are areas whose structure superficially re- 
sembles that found in the cuticle of Epistylis, which yet are 
formed of these independent, but allied fibrillations. I have 
found such in forms nearly allied to Epistylis. Single fibrils 
of this sort when they lie in areas of rather fluid, vesicular 
structure, slip easily in their extension or contraction through 

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the surrounding foam. They are not as a rule bordered by 
true lines of alveoli, and may lie in areas whose inclusions 
are often heterogeneous and irregular in size. They are 
independent of organization in the surrounding structure of 
Biitschli, or other visible structure, because they are them- 
selves the organized contraction areas. By their contraction 
they can bring together separated areas, or cause change of 
shape in the areas they traverse, for they are commonly 
attached at their extremities to some pellicular formation, 
with whose substance they are continuous. They frequently 
become tortuous, or even plicated, in line during contrac- 
tion, a phenomenon seen also in membranes, or in complex 
muscular fibres. 

They may arise suddenly in areas where the interalveolar 
substance has been up to that moment markedly fluid; and 
they may again return, after a variable time of function, to the 
same state ; such metamorphosis being accompanied by those 
optical changes characterising ectosarcal organizations and re- 
distributions of the elements as seen in Biitschli's structure. 

Returning for the moment to striation associated with con- 
tractility as an optical phenomenon of the substance, and 
laying aside the distinction between striae and fibrils, the 
following variations were noted. 

Strial emphasis may involve all, or a part only, of the 
continuous substance, and may or may not involve the walls of 
adjacent alveoli ; therefore a marked striation may be bordered 
by spherical, or nearly spherical, alveoli which, even during 
active contraction of the strial lines, may not change their form. 

Striae in living protoplasm may be in number independent 
of the number of rows of alveoli amongst which they lie. 

Their thickness varies greatly in different instances, as well 
as intermittently, or rhythmically, in the same series from 
moment to moment. Compared optically with the rest of a 
given reticulum at such times, they are like cords or strands 
of a different constitution. Isolation of small portions of 
their length shows them to be a physical differentiation, 
and mechanical experiment shows them to have a definitely 
resistant and often highly viscid consistency. 

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The length of striations varies to any extent, and may 
do so independently of a quite stable linear arrangement of 
vesicles along their course. It will vary in a series or in a 
single stria from moment to moment. Striae may lengthen 
or shorten rhythmically or intermittently. During shortening 
they may become more tenuous, delicate, and finally less viscid 
and refractive, and even disappear from sight or from all but 
the closest scrutiny. Or they may become thinner and more 
obvious optically, because more refractive. 

They may become thicker and more refractive. Those 
striae which I have distinguished above as true fibrils, become 
always thicker and more refractive during shortening contrac- 
tion, and thinner and more refractive in extension. But striae 
may lengthen actually by implication of new substance and 
not by mere displacement of already organized substance. 
They may shorten likewise by relaxation of such organization 
along lines once emphasized. In other words, the physical 
conditions in the finer foam on which they depend may 
be as unstable as similar organizations in the structure of 
Blitschli. Such phenomena as these latter are best seen in 
the development of eggs, where the elements are constantly 
undergoing metamorphoses of structural arrangement. 

Striae may travel from place to place without visible dis- 
turbance of an existing structure of Biitschli, or arrange- 
ment of its elements. In such case, they progress either as 
a wave of influence upon interalveolar substance, or as actual 
(lisplacement of this along interalveolar lines. 

It is common in contractile tissues, or areas, whether stable 
or unstable, among Protozoa, and also Metazoa, such as hydra, 
rotifer, frog, and starfish and sea-urchin embryos and larvae, 
to find ladder-like arrangements of the vesicles, of various 
lengths, extending in a number of directions with respect, 
not to a common centre, but to many centres, and as it were 
growing out of each other at various angles; the optical or 
linear emphasis beginning or ending at any point, with or 
without extension of the alveolar walls of the structure on 
which it is marked out. A kindred incoherence of divergent 
emphases may be seen in the unsegmented egg of starfish and 

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sea-urchin and rotifer eggs before the protoplasm reaches a 
more uniform arrangement preceding first cleavage. In some 
living epidermal cells of a small marine fish, I have found a 
curious whorled effect produced by the interalveolar substance 
lying in somewhat concentric but branched lines, these con- 
trolling the form of the network to such an extent that the 
optical reticulum was in places triangular ; yet closer observa- 
tion showed that the vesicles themselves had true fluid contours 
notwithstanding. As pathological states set in, preceding death 
of the cells, the concentric, or whorled, arrangement gave place 
to a more and more regular and even structure of Butschli, 
the relaxed continuous substance becoming less obvious, the 
vesicles swelling very perceptibly and their hue changing also 
to a darker and bluer tone. This by lamplight. 

In transversely striated muscle bands in a rotifer, I have 
seen coarse striae which were visible in states of medium 
extension, separate during greater extension so as to become 
resolved into double striae. Between these, vertical alveolar 
walls could be seen at happy moments of optical conditions. 
At first only each alternate stria would so separate, but in 
rare and extraordinary extension of the muscle band the other 
alternating striae would half reluctantly separate also, and show 
their structure to be the same. Here were evidently reserve 
powers of extension and of elasticity. 

The rapidity with which the elements of a given vesicular 
structure can be re-organized for contractile activities, is a 
very important point which must not be passed over. If I 
say that such can take place "in the twinkling of an eye" it 
may sound like exaggeration, yet such is indeed often the fact. 
Of course where an existing structure of Butschli remains 
unaltered, the contraction is referable to the ever present 
ectosarcal nature of the continuous substance, having ready 
prepared just the structure for organized action of the sub- 
stance to take place in.^ 

[97] Ci^^ ^^^ ^"^y but external striae, both as regards their 
origin and mode of formation. 

Most remarkable are the phenomena of the aster rays in 

^ See Substance as Such. 

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starfish and echinus eggs, first observed by me in 1893 at 
Wood's Holl, when they formed one of the strongest incentives 
to these researches. 

Biitschli ascribes aster appearances in segmenting eggs to 
an optical effect of linearly placed and stretched alveoli, and 
to illustrate this, has figured a cross section of a preserved 
sea-urchin egg, showing in the "attraction sphere" an indu- 
bitable radial and linear arrangement of the vesicles. While 
an optical striation is undoubtedly produced by the condition 
shown, I am unable to identify this with the appearances of 
the living egg ; and it is surely impossible to explain thus 
other phenomena seen there. In the " attraction sphere " as 
figured, radial increment of substance is compensated for by 
increase of size in the alveoli whose walls form the striae ; in 
the living egg, where the outer cytoplasmic foam diminishes 
the size of its vesicles towards its periphery, such a condition 
ceases to be possible ; moreover the rays do not appear in 
divergent pairs after leaving the "attraction spheres," as, if 
they were due to lines of alveoli, the conditions of increasing 
number and decreasing size of the vesicles with increasing 
diameter of the egg mass would seem physically to require. 

In the living egg the aster rays follow at times very waved, 
or truly curved, courses which are not simply complexes of 
lines formed by alveolar walls, but are irregular with respect 
to even single walls. The rays wander about also in a variety 
of ways as referred to above, often progressing more as filose 
formations than as waves of contraction influence, yet at other 
times gaining, or losing, emphasis in a truly contractile man- 
ner. They have no fixed course ; they curve more or less from 
moment to moment ; they extend themselves now here, now 
there, or conversely shorten ; they even ramify and anastomose 
like true filose formations ; yet, during such changes the sur- 
rounding, or even bordering alveoli, may show no change of 
contour and even no change of place. 

At one point in their history they extend far beyond the 
median line of an as yet undivided cell mass, passing between 
the rays proceeding from the opposite aster, and being in- 
terlaced changefully amongst these. After an innumerable 

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series of changes and much wandering, they become more 
and more restricted in each half the mass to their own side 
of the plane of the coming cleavage. About this time, begins 
an irregular fusing of the tips of aster rays with each other, 
along the course of the coming plane. At first they form by 
this means a crooked sort of network rather parallel to the 
cleavage plane. This becomes more and more straightened 
out in its own plane, and at the same time there is a slight, 
but perceptible, increase of interalveolar stuff along the same 
plane between the alveoli, or, perhaps, merely a spreading 
out there of the substance of the fusing rays. At last there 
is a distinct pellicular, or plate-like, thickening along the 
coming cleavage plane. ^ 

The same process being gone through with on both sides 
the same line, the pellicular plate is double in origin, although 
optically single. On each side of it for a variable time is a 
perfect alveolar layer of vesicles which are unlike the others 
of that central region of the cytoplasm, being small and like 
those of the general periphery ; with this they are, in short, 
continuous in all optical appearance, being truly a preformed 
continuation of it. At this time the rays which spring from 
the "attraction sphere" are very long and in general fused at 
their tips with the general peripheral pellicle. At the same 
time similar thickenings of interalveolar substance are seen, 
tapering from their point of origin as they pass inwards, and 
ending in undifferentiated continuous substance before they 
reach the centrosome. This they never very closely approach 
since the true astral rays maintain precedence upon the alveolar 

Although during actual cleavage the astral rays, or certain 
of them, show transiently rhythmic changes which charac- 
terise true contractile fibres, I do not think that the cleavage 
may be assigned to this alone. For, there is a wave-like 
shrinking apart of the double pellicular plate, which is inde- 
pendent of the rays and which doubtless assists, and could 
accomplish it alone. It is of the same nature as the activity 

^ A somewhat similar plate-like formation was described by Camoy for Nema- 
tode eggs, but of this I knew nothing at time of my observations. 

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by which the endosarc of Vorticella is split apart to form a 
new cleft, and represents a phenomenon common in proto- 
plasmic masses, especially in pellicular membranes, in which 
one frequently sees wave-like modulations. 

At different stages in their ever changing optical emphasis 
and physical conditions, the aster rays show that difference 
of reaction to chemicals which was spoken of under pellicles 
as characteristic of th& substance in its -varying states of vis- 
cidity or of contraction. A few seconds of time may make 
a marked difference in reaction of these structures to harden- 
ing fluids. It should also be marked that a false appearance 
of viscidity, or of such physical states, can be produced by 
too sudden changes in the density of reagents, or by osmotic 
conditions caused by them. There is no doubt but that meth- 
ods which do large justice to finer structural relations of the 
protoplasmic elements, fail often to produce diagrammatic 
karyokinetic figures, and it is therefore peculiarly unfortunate 
that the presence of these structures should be made a test 
of preservation, or methods. On the other hand, such appear- 
ances are frequently had by less careful, more severe, or even 
violent processes. Besides the results of my own special experi- 
ments on this point, I have been struck with those gained by 
others. In one case, during efforts to improve, by re-staining, 
some unsatisfactory material, the investigator had plunged it 
by mistake from ten per cent into ninety per cent alcohol, but 
was rejoiced to find that it gained thereby fine karyokinetic 
figures of the kind sought. The astral and nuclear structures 
were strongly but coarsely marked and seemed destitute of 
all finer protoplasmic relations while the c)rtoplasmic structure 
of Biitschli was much vacuolated and altered. It seems that 
vitiating changes may be wrought even after an initial harden- 
ing and staining. Probably no amount of care and knowledge 
of the conditions to be met can be too great for such experimen- 
tal research, so delicate and susceptible are the protoplasmic 
relations. And especially is this care needed where the irritable 
substance is already in a state of organized contractility. 

Single fibrils in protoplasm, as well as contractile pellicles 
and substance membranes were seen to become waved, or tor- 

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tuous in line during contraction. Just so did those more com- 
plexly organized areas found in the footstalk of Vorticellidae 
and in the motile appendage's^ of all Protozoan organisms. In 
a similar manner were bent and made tortuous such Metazoan 
contractile areas as cilia, and even complex muscle bands, such 
as the head-attachment-muscle-band in rotifers. There seem 
to be conditions in which the line of more evanescent contrac- 
tion waves may be held by the substance for variable periods. 
There are curious structural relations in the muscle of higher 
Metazoa which hint that this mode of obtaining more space for 
mass displacement has been utilised and the protoplasmic differ- 
entiation organized to increase by minute plications retractive 
power of the muscular machinery. Nay more, by using the 
added machinery of jointed endo- and exo-skeletons, such re- 
tractive plication has been still further increased ; and so the 
work of economic extension of energy, on the same basis always 
as that used for the primitive powers of the substance in its 
initial organizations, goes on. 

[98] To sum up the foregoing : I find an enormous number 
of true striations and fibrillations of protoplasm over and above 
those optical and psychological emphases noted by Biitschli. 
I find, moreover, that while the latter, as lamellar films, ex- 
press the arrangement and grouping or extension of alveoli; 
the former must be characterized as products of actual physical 
modification, or increment, or organization, of the continuous 
or interalveolar foam for physiological function. Under this 
head fall all the host of fibrils and filaments as well as all 
striations referable to filose or contractile activities of the 
continuous substance. 

These are therefore to be regarded as substance structures^ 
rather than as basic protoplasmic structure. They are substance 
tissues and substance organs. 


[99^] It must be borne in mind that in discussing contraction 
and its allied phenomena, I speak only of those structures, or 
areas, or masses, of the substance in which the actual changes 

1 The whole section should be read under this number. 

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characteristic of contraction take place. There are always areas 
or masses of substance which share passively in such displace- 
ment, just as fluid alveolar inclusions do. In a Vorticella for 
instance, displacement of the whole body mass is caused by 
contraction phenomena in certain cuticular structures, aided in 
some cases by fibrils running through the body mass. Here 
the statements should be taken as referring to the active areas, 
not to the passive. Even in active areas there is much dis- 
placement of passive material, but this is covered by the terms 
of my statement. 

It is to be regretted that my researches do not throw more 
light on the true nature of the initiative force in that mooted 
property of the substance, — contractility. They seem, indeed, 
rather to render final knowledge of this sort more difficult and 
improbable than it was before they were made. Whereas be- 
fore, the problem rested at a certain structure which could be 
measured and dealt with optically to some extent, and if not 
referable to this, could then be handed over to the physicists 
for molecular hypotheses ; it has now receded, — still associ- 
ated it is true with the same physical structure, — but far 
beyond our optical resources. 

Yet perhaps we are not altogether losers by these new facts, 
for, if, by withdrawing from our reach the elements of the 
problem, they seem to tend to mystification, they bring also 
new assurance that, so far down in the scale as substance and 
substance changes of this sort can be traced, or even inferred, 
we have still the same phenomena to deal with, and these are 
still associated with the same physical facts of a certain struc- 
tural organization of the two sets of protoplasmic elements. 
By so much therefore they have more clearly defined the prob- 
lem to be solved, the difficulties to be met ; making. ready for 
the touch of some master mind unified material without which 
the truth might hardly be grasped. 

As far down as we go, we are still limited to seeing and 
describing the effects of contraction rather than the actual 
contraction phenomena. What we see is always displacement 
of living substance by a wave-like impulse in the direction of 
shortening, the displaced substance, then taking its way along 

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lines between the alveoli of the given vesicular structure, and 
in a direction at right angles to the direction of shortening. 

All my facts point to but one conclusion ; that as far as 
we can follow structure, contractility is not one simple direct 
change in the living substance as a mass, nor even of the con- 
tinuous substance, but that in this latter too it is a complex 
series of phenomena, based upon the physical form of the 
substance as a visco-fluid foam, but nevertheless not directly 
explicable by these conditions. 

Contraction of the living substance as visible with the high- 
est powers is found to be always organized. This organization 
is always upon a basis of foam structure, either of the struc- 
ture of Biitschli, or of a finer froth of the continuous elements 
of this. At any point it directly concerns the interalveolar 

Down to the limit then of microscopical vision, whatever 
portion of a protoplasmic mass is involved, are found those 
familiar phenomena which characterise muscular contraction 
in large masses — as of a frog's leg. That is, there is always 
displacement of mass at right angles to the contraction, causing 
increased diameter at right angles to the direction of shortening 
or compression. But this displacement is seen to be due, not 
alone to mere elastic change of shape of alveoli of Biitschli's 
structure, but primarily to actual displacement of interalveolar 
substance by a wave of contractile influence forcing it onwards 
along lines of least physical resistance. Whether these lines 
of least resistance are relaxed viscosity, or organized paths, or 
merely such as are afforded by the fluid foam structure cannot be 
definitely asserted. They pass at least between the alveoli of the 
existing structure along such lines as their arrangement per- 
mits. Elongation, by compression of the alveoli of the struc- 
ture upon which organization of function has taken place, is 
due both to contraction wave and to interalveolar displacement. 
That the actual lamellae of the alveoli are but passively involved 
in contraction, at least in its maintenance, has been shown. 

These facts remind us again that, at any given moment, the 
true protoplasm — the living, functioning substance — is the 
continuous substance, and that though the inclusions and per- 

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haps the lamellae of Butschli's structure are utilised mechani- 
cally and possibly chemically also in organized contractile 
phenomena, they are not of prime importance. 

Since the living substance wherever examined, down to 
the smallest subdivisions of interalveolar substance in which 
change of shape can be detected, showed such optical and physi- 
cal phenomena as characterised contractile changes in Butschli's 
structure when viewed with equally inadequate powers ; there 
seemed no escape from the conclusion that just so far as this 
must we associate with such phenomena, with such contractile 
manifestations of the substance as such, structural changes like 
those seen in gross on a basis of Butschli's structure* 

New Structural Formula for Protoplasm. 

The facts as they appear to me, seem in mosaic to offer the 
following tentative formula for the living substance. 

[loo^] Protoplasm is a very complex emulsion, having the 
physical arrangement of a very finely subdivided, variably 
viscid, foam, which characters are coextensive with the con- 
tinuous element of all visible optical reticula. 

The substance which at any given moment forms in all sub- 
divisions of the foam the continuous element, varies its vis- 
cidity by some unexplained changes within its finer structure, 
so that from a very fluid state it may almost instantly become 
viscid to varying degrees, even to a semblance of true solidity. 
It is subject to displacement by contraction activity which may 
be rhythmically organized, or may be of a filose nature. Such 
changes may affect all, or a part only, of visible reticula, so as 
to form of the interalveolar substance simple or compound 
fibrils, or threads, or plates, of a different physical and optical 
quality. These having necessarily the course of the continuous 
substance must form optical striae simple or branched, or an 
optical reticulum, the arrangement of whose lines, but not 
whose mass, is governed by mode of distribution of the emulsive 

The continuous substance is at any given moment the physi- 

^ The whole section should be read under the head of this bracketed number. 

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ologically active element of protoplasmic masses. In its last 
resolvable arrangement it has always the form of a pellicle or 
membrane surrounding fluid inclusions. It is the contractile 
and irritable substance. It forms all living contact surfaces. 
It is the sentient substance.^ It is the bearer of those physi- 
ological powers, functions, habits, and instincts which charac- 
terise the living substance. Upon its response in character of 
its powers, or properties, to specific and general environment, 
depend all the physiological phenomena characterising areas, 
masses, or organisms, as such. It is homogeneous throughout 
all areas alike, as to its intrinsic powers and characters, but 
not as to the specific, or habitual, expression of these, which 
varies with its chemical or physical contacts. (See following 

All pathological or abnormal states of the organism, or of 
organs or areas, are directly due to abnormal states of the 
continuous substance, and secondarily to abnormal difference 
in the specific or immediate stimuli for this; either in the 
affected area or in those which control, or are controlled by, it. 
Death of the organism or the area is due to irretrievable dis- 
organization of the continuous substance in the chief tributary 
areas or in the whole area. The initial stages of decomposition 
are always the dissolution of an organized arrangement of the 
continuous substance, and, after variable protoplastic activities 
of this, a dissolution of its organized finer structure, and the 
replacing of both organizations by a purely physical vesiculation 
having an optical appearance of a fine structure of Butschli.* 

The discontinuous elements, or protoplasmic inclusions, are 
most heterogeneous, in their ultimate subdivisions defying 
chemical analysis, as they do all effort to separate them optically 
from the lamellae of the continuous element as such. 

They form the chemical contacts or specific controls, that 
is, the specific environmenty of the living substance. They are 
the secretions, assimilation products, excretions, and hoarded 
reserves of the living substance proper. In Butschli's structure 

^ See following sections. 

' This statement is the outcome of many direct experiments with areas of living 
organisms and whol^ organisms. 

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they seem in general to play a passive mechanical or physical 
r6Ie and are strongly hinted by the phenomena to be for the 
most part, and for most times, segregated excess, waste, or 
reserve material of the living substance. 

By holding the place of continuous element in the proto- 
plasmic foam, the living substance has its contact, or more 
strictly speaking, its contacts, almost infinitely extended and 
multiplied. For every pellicle is a contact surface of the 
irritable substance as such, and a possible contractile membrane; 
and to this every alveolar inclusion, as well as every local dif- 
ference or activity within actual touch of it, bears as truly a 
relation of stimulus or control, as does external environment to 
the substance as organism. 

To the irritable substance, contact is stimulus : to the living 
substance, contact is environment. 

Heretofore in biology, environment has been used in rather 
a limited sense for the sum of influences which affect an organ- 
ism from without, as opposed to its own intrinsic forces; there- 
fore as modifier of its inherent tendencies as organism, and chief 
factor in determining the final result of organization. 

From the new standpoint of the substance as such, which it 
is my privilege to preach, biological use of the term environment 
must be widened to give it its full value canying our thought 
of it into the minutest subdivisions of living substance. 

It is at once plain what a magnificent device is the physical 
form of the substance for multiplying and extending contacts ; 
and, given the physical variant of viscidity, as well for preserv- 
ing, conserving, and unifying, stimuli ; for segregating incon- 
venient and inutile substances ; and for providing at any point 
a homogeneous basis for organized activities. As by convolu- 
tions in the brain, thought surface has been, and may yet be, 
greatly extended; so by structural subdivisions of a fluid foam, 
the environment of the living substance can be extended and 
multiplied until it may even be coextensive with a primal 
irritable stuff. 

For what is at first but a minute and momentary contact for 
the substance as mass or organism, can presently be transformed 
into contact for the substance as such, whic^ is now limited 

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optically by subdivisions of a visible structure, but which the 
known subdivisions of this again extend, and which the phe- 
nomena of the invisible substance prove to have no limit assign- 
able by us. A concrete instance will help to an understanding 
of this broad statement. 

When an amoeba, responsive to the touch of a passing organ- 
ism, engulfs it, the protoplasm, obedient to its physical form, 
surrounds as a pellicle the ingested mass, and thus that first 
slight point of external touch becomes at once extended into 
an internal contact many hundred, or thousand, times its first 
area. As digestion proceeds, the chemically altered food be- 
comes more or less a watery solution and in this form is grad- 
ually diffused throughout the whole organism until, both by 
this and by wandering of interalveolar material and of vesicles 
of Butschli's structure, the living substance may to its farthest 
morsel suffer change of environment, which may be by way 
of extension, or renovation, or actual change, of stimuli at 
multiple contact surfaces formed by lamellar films. 

Thus by ingestion of food is the specific environment of the 
substance not only extended but perennially renewed. Nor do 
the phenomena differ in any essential in those higher organisms 
furnished with most stable and complex machinery for the dis- 
tribution as well as ingestion of food. These facts hint that 
there is a more extended significance than has been understood 
in the habit of reducing food substances to a diffusible form, 
for they are now seen to be thus prepared not for the stomach 
membrane only, but for the universal physical form of the 
living substance. 

The subdivisibility of the foam structure, the general physi- 
cal plasticity of such a form, plays an important r61e in arrange- 
ment, and in procuration also, of specific, internal environments 
for the living contact surfaces or vesicular films. For, as was 
shown above, relatively large quantities of inclusion substances, 
as well as of living substance, can be transported through, or 
amongst, very stably organized areas of physiological function 
to other parts of the mass, or even outside it. 

From the physical form of the substance results great 
economy too of the vital stuff. Growth of areas, masses, or 

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organisms can be effected to an enormous degree by mere 
accretion of passive, non-living material, without the actual 
living stuff being increased at all, for there is a very wide 
range to the thinness of alveolar films in each structural series. 
The amount of true living material, as we know it even, is 
probably very small in any organism. In the young organism, 
notably in *the developing egg, the network substance is often 
markedly thicker than it is in later stages. In the minnow's 
egg and in the starfish egg this was beautifully shown. While 
this may not be a rule it is true of many cases. Again, one 
adult organism may be larger than another and still hold actu- 
ally much less of the living substance. 

From the physical form of the substance, again, arises 
infinite opportunity for contractile service of an organized sort. 
Of physical necessity, the very contact between two vesicles 
produces planes and lines of continuous and direct field for 
organized contractility, — nay almost compels it, if the irrita- 
ble and contractile nature of the lamellar substance be taken 
into account. 

For receiving and transmitting physically all physical impacts, 
as well as for appropriating or reacting to, chemical contacts, 
it would be difficult to conceive of any combination more per- 
fect than that afforded by the multiplied and extended lamellar 
membranes of widely variable viscosity and tenuity and curva- 
ture which the facts have seemed to express as the form of the 
living, irritable, contractile, sentient, substance. 

The Living Substance; as Such; and as Organism. 

As indicated in areal differentiation, the problems confront- 
ing us in organization of the foam elements in their specific 
relation to each other, are but those with which we must deal 
in understanding the origin of physiological organs or areas, 
not excepting those of the highest organisms. 

[loi] Conversely, it is even more true that the problems 
confronting us in the most complex and stable organs of the 
higher animals are identical with the problems found in the 
living substance as such, to its smallest visible subdivisions. 
Nor does it end here, for there is indubitable evidence of 

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protoplasmic extensions invisible to our strongest magnification, 
having still the same structure, the same characteristic dis- 
placements, which would yield to bettered tools the same 
optical phenomena as belong to large organized masses, — such, 
in short, as pertain to the characteristic physiological activities 
of protoplasm. 

[102] Biitschli found that mass has much to do with the rela- 
tive character and duration of phenomena in his artificial foams. 
The living substance seems to transcend such physical limita- 
tions. That these statements are not merely wild speculation 
or inference may be shown anew by a single instance, and it 
is thought that the filose phenomena cited previously also bear 
them out. Many others could be brought in evidence but for 
lack of space. 

The cup-shaped film, or "collar " of many Choano-Flagellata 
is so tenuous in greatest extension that when magnified eight 
thousand diameters, though it may be visible obliquely and later- 
ally, one cannot see it in direct, transverse, optical extension, 
not even as a hair line ; yet it is even then thicker than many 
webs and veils Gromia extends. 

By addition of pigment to the water a double set of currents 
can be demonstrated in this film. One current flows up and 
out from the body over the outside of the collar to its edge. 
Turning this, it then flows down the inside, back towards the 
body substance with which it mingles. Small organisms brought 
against the outside of the collar by rotary beat of the single 
projecting flagellum that springs centrally from the anterior 
end of the body, are carried along to the body substance which 
engulfs them in usual protoplast fashion.* At times the collar 
assumes considerable rigidity and the flow ceases for many 
moments. I find that if a jar be given the cover glass, the fluid, 
flowing film instantly contracts, taking the form of a truncated 
cone. During this contraction, striae appear which run from 
the top downwards, gradually thinning, fanning out, and fading 
away as they near the base of the cone. Conversely, they are 

1 These phenomena were first described by Saville Kent whose faithful, patient 
researches among minute organisms have yet to receive their full meed of appre- 
ciation from biologists. In my own researches I have ever found Kent's work 
preeminently accurate and fine. 

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more marked and refractive as they near the edge, where they 
are so closely set together as to appear fused into a thick, 
highly refractive rim. At this time the collar has perceptible 
and constant viscidity, amounting to rigidity, for it bends not 
at all to sharp pressure from other larger organisms, as from 
the appendages of Stylonichia moving about it. 

From the standpoint of a physical and mechanical hypothesis, 
what marvellous and irrevocable transformations of substance 
must instantaneously have taken place in response to that 
passing jar of the cover glass! 

Yet after a few moments' quiet the striae begin to be less 
distinct and the shape of the collar shows a new change. It 
opens now at the top, widening gradually until it again equals 
its first expansion. As it nears this limit, another set of striae, 
but far more delicate, appears encircling the collar, that is, 
transversely to the first set which has by then quite disappeared. 
Finally all striation vanishes, and by their renewed movement 
up and down over the surface of the film the pigment granules 
show that the former fluid state is again active. That it is truly 
a fluid state is shown by the whole film being at other moments 
returned by flow into the body and then again extruded in any 
one of a number of fantastic ways. It may appear next as an 
amoeboid mass extending lobose processes, or beyond these filose 
threads, or as a huge bubble of protoplasm such as Vorticellidae 
are found blowing, the walls of which may here be thick or 
thin, smooth or variably lumpy. This will thin out more and 
more at the uppermost convexity until it seems to burst like a 
bubble. After this the whole expands itself gradually, — the 
irregular protoplasm returning into the body, — and the normal 
appearance of the " collar " is restored. The flagellum may be 
reproduced from the centre of the collar included area by an 
irregular pseudopodial process which undergoes swift recon- 
struction until the normal flagellum is again actively functioning. 

The collar will again later suffer metamorphosis into some 
new mask for the substance. How unimportant here does the 
form of the substance seem in comparison with its habit ! 

If a cross section of a collar film could be made, equalling in 
depth the thickness of the film, we should have a filose-like 

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mass, invisible except when in a contracted state, yet wide 
enough for a double set of substance currents to flow upon 
each other, and possibly with even a third supporting area 
between them, such as obtains in many filose extensions of 
protoplasm; wide enough it certainly proves itself to contain 
the substance and much structure, and rjaom enough for such 
structural organizations and changes as are associated with 
intermittent organized contraction. From the outer surface of 
the cup were sent off at times delicate filose processes whose 
mass being great compared with that of the film looked quite 
large. These have been taken by some observers to be ad- 
herent bacteria. 

[103] In Gromia the contraction phenomena of threads and 
webs are even more marvellous. Such facts, with others pre- 
viously cited for filose activities, go far to convince one that to 
limit protoplasm and protoplasmic phenomena and protoplasmic 
physiological structure to those grosser masses we now discern 
would be to stultify ourselves and to lose many chances for 
valuable effort. If Biology is to be the guide of physiological, 
medical, and hygienic science, which would seem to be her 
highest function, it is such minutise of life history and possi- 
bilities of the living substance, which offer best fruit to that 
willing patience, that single-hearted, loving receptiveness of 
research for which human progress ever waits and upon which 
it still must lean. 

One cannot but wonder with a child-like wonder, at the 
infinite smallness possible to the living substance as such, and 
even as organism, but to refuse credence because of this unimagi- 
nableness would be as childish as for one who had never looked 
through a microscope to disbelieve in the Protozoa altogether; 
as childish as to refuse to believe in those hosts of mighty 
stars which for the naked eye have no being. 

The nature of a physical foam such as postulated here, makes 
possible for protoplasm a disposition of its elements in such 
minute sub-divisions as water has in air; makes it possible for 
the substance as such when seen from afar, — as with our poor 
optical powers we still must see it, — to look as still and struc- 
tureless as those cloud masses which seem to hang motionless 

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for long days upon a mountain peak, but which when climbed 
to and met in close encounter, are found to be hurtling masses 
of vapour, rushing powerfully against the crag on one side and 
melting away like an elusive memory as they strike a wall of 
warm air just beyond, while every foot, every inch, of those 
square miles of vapour is stir and changeful difference. 

[104] The marvels of filose phenomena have been dwelt on 
at length. It is not undue emphasis has been given them 
for they are by far the most widespread and most characteristic 
phenomena of the substance, if we except contractility. 

In ordinary amoeboid movement there are two sets of phe- 
nomena to be reckoned with; a purely passive flow governed 
by impetus received from local substance contractions; and a 
more mingled flow in which the same cause lasts longer and 
passes as a wave of contraction along certain lines of the sub- 
stance causing more or less steady and continuous displacement 
before it of the substance in its path — like that of interalveolar 
substance along the fibrils, or as in filose phenomena.^ 

In amoeboid displays the structure of Biitschli or a coarser 
is largely implicated, the finer foam of the continuous sub- 
stance being of course involved. In true filose activities the 
displacement is limited to the finer froth. This expresses I 
think, the whole ground of difference between them. I do 
not deny to protoplasm motion of the sort which Biitschli 

^ Having spent hundreds of hours in watching the flow of amoebae as well as 
of many other and diverse Protoplasta both lobose and filose, besides Myxomycetes; 
I venture thus to express myself counter to certain widely accepted opinions. The 
phenomena have been looked at too much from a standpoint which accepts the 
most obvious phenomena as of prime im]>ortance, using these for the starting point 
of observation; and which is further baffled by that most treacherous of all rela- 
tions, a time, or sequence, relation, ZTgrnng post hoc ergo propter hoc. The subtlety, 
complexity, and fleetingness, of protoplastic phenomena at their simplest, together 
with the fact that at any given moment many causes and many effects are crossing 
each other as impulses and impulsions of fluid and readily displaced substance ; 
have seemed to me to make the study of amoeboid phenomena quite other than 
one could imagine from any printed description to be met with. One might easily 
give a lifetime to unravelling these phenomena and without much result if one 
followed the established precedent in this line, that is, looking for and seizing upon 
change of external contour, and grosser internal and external displacements and 
correlating these which, as a matter of fact, apart from mere mechanical displace- 
ment caused at times by impetus of one or another, have usually little or no vital 
relation to each other. 

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demonstrated in his physical foams, for these researches proved 
nothing for me more conclusively than that the physical op- 
portunities of its form are made large use of by the living 
substance. But this is mere incident in its whole life history. 

[105] All the facts relating to the filose habit of the substance 
are most interesting when taken in relation to the possibility 
that Metazqa had their rise not from an ancestral amoeboid 
stem but from an ancestral rhizome of Filosa. The new facts 
seem to suggest that the Metazoa arose rather from a filose 
type which by coalescence, or areal differentiation, built itself 
up into compound masses. If by coalescence, the substance as 
such showed respect to that position in the mass in which it 
newly found itself, exactly as in each individual it had through 
all its ceaseless flux respected its relative position; for it must 
not be forgotten that in these protoplasts the substance as such 
is ever changing its position in the mass or organism. 

By such hypothesis, complexity of organisms would arise 
by extension and multiplication of areal differentiations which 
were but rearrangements and redistributions of the two sets of 
foam elements in relation to each other; in short by mere con- 
tinuance and extension of exactly the same state of things that 
existed in the individual as a starting point. 

This hypothesis would be a true description of the phe- 
nomena which can be seen to take place in development of a 
starfish, sea-urchin, rotifer, annelid, frog, or hen's egg. 

It is a very difficult thing to grasp this idea of a sensitiveness 
of the substance as such to its position in the mass; it cer- 
tainly would not occur to one ci priori. The reader is reminded 
of the viscosity rhythms of the substance in sea-urchin and 
starfish as correlated with its position in the whole mass 
whether this were of single cells or of many cells. In all pro- 
toplasts the same thing is seen. In the phenomena of leuco- 
cytes of crab's blood it seems that organization of this sort is 
instinctive among even wandering cells of a Metazoan organism, 
under even novel stimulus of opportunity or external conditions. 

A still clearer understanding of what is here meant can be 
given by citation of an organism which, before the heart of 
these researches was reached, was thought to be from the cell 

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standpoint a missing link between Protozoa and Metazoa. 
From the standpoint formulated by my facts, from the stand- 
point of the substance, it is now seen to be even more valuable 
as a link between these two, and as a most expressive type of 
substance life writ larger than common. 

There is a Heliozoan described by Leidy and others, Raphi- 
diophrys elegans^ which lives at times as isolated individuals, and 
then again, for variably long or short periods, in coalescence 
with others to form a wandering colony. 

As single individuals, the animal does not differ from all 
typical Heliozoa. Its filose pseudopodia are used for locomo- 
tion, for prehension of food, for tactile purposes, — for what 
else we simply do not know. In its isolated state, these proc- 
esses are freely produced on all sides of the periphery, and are 
on all sides alike in kind as in origin and function. 

In the coalesced, or compound, state, the individuals form a 
colony whose units are separated from, as they are joined to, 
each other by bands or bridge-like extensions of protoplasm 
which are formed by modification of the usual processes and 
make living links between the animals. Between these bands 
the water surrounds the colony ever)rwhere. 

The remarkable thing is that no matter what may be the 
number of individuals so coalesced, and Leidy states colonies 
contain at times upwards of thirty (I have not seen so many), 
the usual filose pseudopodia are formed only at the periphery 
of the whole mass. That is, each Raphidiophrys lays aside, 
for the term of its union with the others, the habit of filose 
formation at all those portions of its mass, which, though 
peripheral still to itself, are not peripheral to the colony as 
such. From those portions of each unit, which, through coa- 
lescence, become in a sense interior to the colonial mass, there 
are produced only bands or bridges of thick protoplasm, both 
ectosarc and endosarc in their usual relations to each other, 
and these form the ties between the individuals. Those units 
which are wholly internal to the colonial mass, produce no 
filose processes but only connecting bands. 

Two points of utmost weight must be given due emphasis in 
studying this physiological marvel. First, that the substance 

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of each individual is in ceaseless change of position with respect 
to the mass of that unit; and further, with respect to the mass 
of the colonial organism as such; besides which, there is the 
ceaseless interchange of substance and function between ecto- 
sarc and endosarc, and finally a ceaseless interchange of sub- 
stance between the units. The units are covered throughout, 
over their whole periphery, by the usual ectosarcal formation. 

[106] For the substance as such, then, there is no quiet nor 
any persistence of position. Yet each portion of that substance, 
wherever it passes or pauses, knows where it is and how to 
play its part as citizen-substance of the coalition. Surely this 
is more marvellous than the imagined stable diversity and 
interaction of Metazoan cells, if indeed we still dare to say that 
in them is greater actual repose of the substance as such 
beneath the mask of form of organ and organism. 

The final touch to wonder is given when it is seen that the 
units of this coalition are, as organisms, surrounded at all points 
exactly as usual by their normal external environment, for at 
first they are joined by their filose processes between and 
around which, as later with respect to the bands, the water 
freely passes. 

If, in the starfish egg, the phenomena are such as to incline 
one, if one does not bow to the mosaic hypothesis, to think 
there must be in the egg some coordinating area or factor 
which it, as organism, possesses, — a sort of brain direction 
owned by all the cells in common, — how can one bring such 
an interpretation into play here, where the units are not parts 
of a single mass formed at one time by one parent, nor even 
indirect products of two which peculiarly dovetail their prop- 
erties and "ids "; but is a coalition of stranger organisms. In 
which should one say the control resides } Or shall we think 
one unit perhaps assumes entire direction while the rest hold 
in abeyance their control power as they hold part of their 
pseudopodial formation.? Then how is this common harmony 
of action got at? And how reconcile it with the retention in 
part of control by the units.? For it is hard, too hard indeed 
for me, to believe that the single controlling centre would be 
able to manage this feat. Finally, how can the units as such 

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be in any case under such control from one centre when the 
substance of the whole is in ceaseless flux from one to the 
other of the whole group and in all parts of the units. 

[107] The control in this case must, it seems to me, be in- 
trinsic to the substance as such^ and not to the substance as 
organism. I think, in short, that one must reasonably believe 
that the living substance, as far as it can be traced optically, 
is not only irritable and contractile substance, but that it is 
already conscious organism. By conscious I mean that more 
basic form of it which I have above termed sentience. And I 
think that the phenomena recited already in this paper will of 
themselves alone bear out to great extent such a conclusion. 
Sentient, but subject to such direction from organized centres 
of this same sentience as may be supplied by the nucleus, and 
other areas of organized differentiation, as the cytoplasmic 
granules, — I assert the living substance to be. 

The complacency with which the facts on all sides of my 
investigations lend themselves to a founding of the Metazoa 
on an ancestral rhizome of Filosa, is reflected in these further 
characteristics of the Filosa. The Filosa, as I should widen 
the use of that term, are by far the most numerous and widely 
distributed so far as we know, for they would include all marine 
Heliozoa, and the Radiolaria as well as the Foraminifera, in- 
cluding Gromia and similar forms. As a group they are 
marked by a tendency to coalesce ; and to multiply the nuclei 
without actual separation of nucleated areas. As a group they 
are marked by a plasticity of organization, and show a tend- 
ency to areal differentiation of structure and a power of adap- 
tation to general environmental conditions in a number of 
ways, notably by formation of free-swimming, flagellate, larval, 
masses. Their differentiation is of the special character which 
I And to distinguish animal organization ; that is, it tends to 
emphasis and intensification of irritable and contractile function 
by organization of the elements upon an ectosarcal basis. 

As a class the Filosa are marked by habits of substance 
metamorphosis for purposes of perpetuation, such as formation 
of free-swimming, flagellate larvae, and also of a minute germ- 
like posterity which are freely motile and suggest sperm. 

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Besides this, they bud, and multiply by fission also in many 

As a class they have a habit of, or tendency to, actually con- 
jugate, which must not be confused with the mere coalition 
phenomena, for in this former case the nuclei of the individuals, 
which may be two or more, are diffused for the time and then 
again reorganized. These actions result after variable periods, 
it has been observed, in reproductive phenomena, and so the 
Filosa are open to the increased plasticity of destiny which 
such a mingling of individual masses or substance seems to 
produce in the substance within fixed limits. 

[108] To sum up the above remarks, my mind is drawn with 
some force by the suitability in all ways of the filose protoplasts, 
in habit as well as in form, for the evolution of a race of com- 
pound and complexly differentiated substance masses, with 
multiplied latent possibilities, all parts of which seem to retain 
to greater or less extent the spinning habit. 

True Biological Standpoint. 

[109] But after all, whether we consider past forms as the 
root of present forms, or these by themselves, the thing of 
greatest moment to us is neither form nor structure in linked 
gradation from age to age, but rather that of which all form, 
all structure, are but the casting, the mould, the framework, 
and the mask. It is the living substance as such, its fleeting 
activities and their meaning which should concern us most. 

[no] Surely all the facts urge a somewhat changed stand- 
point in biological research. We are not denied an ultimate 
return to purely physical interpretations. These are not yet 
pronounced impossible nor even improbable; but we are bidden 
for the time to a physiological standpoint as more immediately 
fruitful. That there is a physical machine seems now more 
than ever certain, but that the machine within our present ken 
is the wonder-working machine, seems less than ever predicable. 
We are forced to see that the final automaton, if such there 
be, must be allowed to be, within an incredibly minute mass, 
almost infinitely self adaptive to innumerable contingencies. 

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The final problem falls thus, for the time, wholly within the 
realm of metaphysics, and the biologist must become again 
the naturalist if he is to win from his toil anything but 
negative results. 

[ill] The facts seem to me to instruct us that, to the liv- 
ing substance as such, form, as well as size, is mere incident. 
Structure, so far as it has been detected or inferred, becomes 
for the time being secondary in importance, as it is in origin, 
to the habit of the substance; and we are warned that when 
regard is had chiefly to visible structure as such, or when this 
is taken to be an explanation of phenomena rather than an 
expression of substance habit; when it is taken tq be the cause 
rather than the effect; the record becomes to us a cypher 
instead of language. 

[112] For structure, as far as I have traced it, was still seen 
to be a mask behind which the most important business of the 
living world is carried on undetected. 

Nature might well be likened to a great spider, spinning and 
spinning the living stuff and weaving it into tapestries; and 
still hiding herself and the ever-lengthening thread of vital 
phenomena behind the web already spun. To nature, the fact 
of prime importance seems to be the substance, — the substance 
— the substance; whether as mass, area, organ, or organism. 
Nothing seems to be of great account with her compared with 
the character and possibilities of the material she is dealing 
with, — and truly it is a most plastic stuff. 

One realises in thinking over such facts as have been here 
described, that, from this standpoint, man's classifications, 
though significant, are not inviolable. He has concerned him- 
self chiefly with individuals, organisms, species, and races; 
with form, and with structure as the exponent of form. 

But it is to the living substance as such that all nature's 
pliant cherishing has regard. Form and structure are used to 
this end; they are not in themselves ends for her. From such 
a standpoint, distinction between individual organisms and the 
substance as such is mere child's play. That has been man's 
game with the substance which he has found strewn about him 
in pieces he can grasp, arrange in lines and group as he will ; — 

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as a child might arrange the pieces of a puzzle whose total sig- 
nificance is beyond him; wondering to find upon so many- 
isolated pieces the same bright colours; trying by these to 
make connection, but little dreaming that the whole is built 
up on a system of grand but subtle lines to which the colour, 
although limited here and there by them, is mere incident. 

From this standpoint, animals, even man the animal's lord, 
is but a piece of the sacred substance, mingled with an enor- 
mous proportion of alien and valueless material. It is nothing 
to nature then what happens to this or that individual, species 
family, or race even. Before these fall like a leaf from their 
place, some most precious substance has already been massed 
under her guidance in other storehouses, or is on its way to 
yet more secure keeping; or the triumphant substance in other 
forms proved better fitted to the new conditions. Again and 
again the facts repeat that form is little or nothing except as a 
means to this end. Whatever form will best preserve the 
living stuff, whatever modifications it can best be cherished 
by, are seized on. As conditions change, one disguise after 
another is assumed by the substance as such that it may run 
the gauntlet of adverse, or hostile, conditions. 

Transmigration is thus seen to be a strictly biological truth. 
In this, too, is rooted nature's apparent cruelty. Creatures 
destroy each other, but nothing is lost; the substance is rather 
actually strengthened. Barring the outcome for the substance 
as such, it is all one to nature whether the bacterium or the 
Buddha win.^ The living substance is free to determine its 
own fate, so only that that substance, stronger and subtler and 
more powerful to control its own immediate conditions survive. 
It is as though the multiplication of forms and individuals were 
but a device by means of which the substance as such should 
be strengthened, — as though a man were to practise one hand 
against the other to increase the strength of either .^ The indi- 
vidual is part of the game, and the conscious or semi-conscious 

1 The ideas of nature and of a deity are not to be confused here, for " God is a 
Spirit " and something, I conceive, quite apart from the order of physiological 
nature as dealt with here. 

' I use device always in the sense in which a mechanician uses it. 

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antagonism of the individual to that truth is merely a useful 
factor in the fact. (See Fosterhood and Heredity.) 

[113] Degeneration, or simplification of form and structure, 
are often as useful to the substance as increased complexity or 
emphasis of function. Its business is to use or to evade its 
environmental conditions. Nature busies herself ceaselessly 
with and for the living substance as such. Transplanting, 
grafting, pruning, and fostering she still keeps it fresh, young 
and unwearied. The separation of one portion of it from 
another in those masses which we call individuals means some- 
thing quite different to her. To her the living substance is 
everywhere continuous. The strange, inseparable, duplex or 
triplex relation of parent and offspring substance is but a 
single though strong hint of her attitude in this respect. 
(See Fosterhood.) 

[114] Only when the biologist knows his substance directly 
in living states can he learn it argumentatively from preserved 
material in cases where the living states are perforce hidden 
from him. He must choose to know it as a living substance 
rather than as dead coral reefs of structure. He must study it 
as activity rather than as product ; — as a ceaseless becomings 
rather than as an achieved and rigid fact. Chief of all that 
these researches hope to accomplish is to tempt the biologist 
from the artificial, the dogmatic, the systematic, mode of 
study; to tempt him to observe the living substance in its 
haunts of structure, in the same way that a naturalist watches 
the habit of the organism as such ; to tempt him to a pursuit 
of substance life, substance structure, and above all substance 
habit. These he will surely find so absorbing that his micro- 
tome and paraffine bath will rust to uselessness on his shelf 
before he next turns to take them down, — after such patient 
devotion of seeing what is to be seen without them, that 
he will to some extent know what he is looking for by their 

If the biologist, in such a naturalist-like pursuit of the phe- 
nomena of the substance will cultivate the narve rather than the 
learned standpoint ; if he will isolate and perfect so far as pos- 
sible his animal faculties and perceptions, such as wild animals 

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use in their daily life, and such as conjurors make use of in 
their sleight of hand and second-sight tricks; he will gain for 
some time to come more knowledge of the living substance as 
such than if he applied the whole weight of historical and con- 
ceptive training to bear on the difficulties. There is to be 
seen amongst expert manual labour and the expert knowledge 
gained in the mercantile business of the world, a far higher 
and more delicate cultivation of the perceptions necessary for 
just this work than is to be found among scientific men who 
have spent years in so-called laboratory training. 

If after his perfecting, so far as possible, these animal 
faculties the biologist finds himself seemingly as far as ever 
from the goal of perfect grasp of the ways and expressions of 
the living substance as such; he has only to recall how difficult 
it is for him to follow the gross movements of whole and large 
organisms across large spaces, — the utter baffling of his keen- 
est attention which is possible to human hands, arms, or even 
a whole body. Thus he will regain a large patience with his 
defeat and be content to serve seven years to learn more of 
the movements and habits of the living substance as such. 
He will then learn how strong, how yielding; how bond, how 
free; that substance is; how seemingly wayward, and yet to 
the limit of our opportunities of knowing it, an expression of 
disciplined and unified results, sympathetic to the sum of past 
experiences, — prophetic of the sum of life yet unlived. 

[lis] To concern oneself much with the living substance as 
such; with the habit of the substance, rather than its form; to 
see, to understand, how the substance expresses itself in rela- 
tion to its environment, both general and specific; to follow it 
in its complex action and reaction ; to learn how it uses, rejects, 
transcends, or passively protects itself against external condi- 
tions; and how it bends itself before these in order to conquer 
by a magnificent "jiujutsu": this seems to me the true bio- 
logical standpoint, for it is the standpoint of the living substance 
as such, — it is the working standpoint of nature. 

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Selection op Environment by the Living Substance. 

I have shown that throughout the Metazoan mass, as through- 
out the Protozoan, physiological function is correlated with areal 
organization of the elements of the emulsive foam. In the 
former as in the latter, organs are areas of vesicular organiza- 
tion ; and in either, all such areas however fleeting are true 
substance organs. 

In the higher forms, there is a more fixed and definite habit of 
structural arrangement, certain relative localities being fixed for 
grouping certain discontinuous elements in a certain way ; while 
in the lower forms, locality for any given mode of organization 
may be as unstable within certain limits as the substance itself. 

[ii6] Nothing new by way of minute structure visible with 
the microscope was seen in the most complex of the series of 
organisms examined ; but merely a more extended and multiple 
and stable organization of the foam elements, and of the habit 
of the substance in correlation with these things ; the contact 
relations thus brought about resulting in quantitative and quali- 
tative emphasis of physiological function, and a more extended 
control by the substance as organism of its general environ- 
ment as a source of supply, or control of its own use of supplies 
obtained from that source. 

[117] We are so used to limit our notions of environment of 
living beings to the external world, and to ignore that which 
forms for the substance a large internal world of contacts and 
opportunities ; so used to think of environment as in a way 
dissociated from — nay, even antithetical to — organisms, that 
it may be difficult, perhaps, to think all at once of external 
environment as a mass of heterogeneous conditions bearing to 
the external contact surface of organisms the same relation 
that the blood of a frog, for instance, or the contents of a food 
sac of amoeba, or even of an alveolus of a structure of Butschli, 
bears to the substance surrounding it. The ancient warfare of 
the race with environment hinders us, mdeed, in grasping the 
truth of these relations and realizing that external environment 
is to the substance as organism akin to the sum of internal en- 
vironment for the substance as such. 

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Living organisms are surrounded as masses by a general 
external pellicle of protoplasmic membrane, within and continu- 
ous with which, is a complex web of delicate, irritable, contrac- 
tile, membranes. The mass pellicle, whether it define the 
limit of one or of innumerable cells, expresses the contact sur- 
face of the organism as such in its relation with environment. 
For it, environment is of a twofold nature, being compounded 
of an internal and an external set of contact conditions. 

The external set of conditions is, to great degree, beyond 
control of the substance, except indirectly through aid of its 
internal environment which bears a constant relation in both 
kind and arrangement to the needs of the physiological powers 
of the substance, and seems to be controlled by it in a twofold 

The internal web of protoplasm, on the other hand, is physi- 
cally surrounded by, and in contact with, an environment upon 
which it acts and to which it reacts in a multiplicity of ways. 
This is more or less completely within its control, yet influ- 
ences it largely and even to some extent controls it ; physically 
and chemically. Being met here by the undying question of 
priority of control, it is possible to say only, that, granting a 
possibly complete, final control of the substance by physical 
and chemical conditions, yet in the existing state of things 
visible to us as structure, the substance is seen acting through 
and upon an existing internal environment. In this way the 
certain conditions necessary for its renewal are detected, pur- 
sued and chosen by the organism. These the living substance 
as such, acting still through existing internal conditions, pro- 
ceeds to transmute and to arrange with reference to its own 
general and local needs. 

[118] The internal environment of the living substance is 
finally, as has been shown, inseparable for us from lamellar 
subdivisions of the interalveolar foam. Even supposing we 
could isolate and retain in a living condition for this purpose 
considerable quantities of the continuous substance of Biit- 
schli's structure, it would be impossible by the most minute 
chemical analysis to determine more of even constant results — 
such as proteids — than that they were either necessary constitu- 

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ents of the living substance ; or constant products of its activi* 
ties ; or a necessary and invariable part of its most intimate 
conditioning.^ And whatever may be the destiny of the mate- 
rials introduced into, or formed by chemical activities within, the 
living substance, their first, inescapable, function is assuredly 
as pellicular or lamellar environment. 

Such physical and chemical conditions as are introduced into 
protoplasm yield but heterogeneous stimuli — a most mixed 
environment — for the substance. Since for organized activity, 
organized specific environment, is needed and provided, the 
bringing together, the chemical preparation, and the peculiar 
grouping, of the various substances which serve in this capacity, 
form the problem to be solved. 

[119] In non-living, or perfectly inert foams, there is a tend- 
ency for vesicles of similar size and bearing similar contents 
to become gradually grouped together, and for these latter 
where fluid to coalesce. It is not such mechanical aggrega- 
tions as these we must explain in protoplasm, but such aggre- 
gations and such segregations as take place under conditions 
seeming to contradict those demanded by the given physical 
explanation. We must account often for a subdivision or 
reduction in size of these kindred inclusions when brought 
together, and for specific redistributions of the continuous sub- 
stance : We must explain complex deportations of substances 
from areas where physiological activities of a sort to which 
these are physically an impediment appear later ; and^ in other 
areas, a rapid massing together of inclusions peculiarly favour- 
able or necessary to the physiological functions called for there. 
Most difficult of all, we must account for these physically 
complex and highly organized phenomena taking place almost 
instantaneously as response, not only to specific chemical or 
physical environment of an accustomed sort, but to a wide 
range of such stimuli, even those never before experienced, 
and to very local stimuli, of a purely physical nature applied to 
some distant area of the organism ; the physiological response 
being often made in defiance of certain strong physical handi- 
caps in environmental conditions. 

1 Finding none suitable, I am forced to create this nonn for a term of my results. 

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[i20] Though finally inexplicable, the phenomena maybe 
formulated in a way which at least unifies the difficulties and 
places the problem in a form more easily grasped and handled, 
when it, like all the radical questions already presented, is seen 
to be a question that pertains to the substance as such. Like 
them, it is but writ large in the organism ; like them, it may 
also be tracked to the ultimate visible subdivisions of the sub- 
stance and found to exist there unchanged in its nature. It 
may be called the selective power of the living substance with 
regard to its own environment, both general and specific. 

Turn where we will in the kingdom of living things, the 
substance in all its complex phenomena, in all its manifold 
forms and phases is still seen to be in act of either pursuing, 
securing, disposing of, transmuting, or using as stimuli, its 
peculiar internal environmental conditions, — the latter phe- 
nomena being but varied aspects of the former. From the 
amoeba to man, it is certain that the substance as organism 
selects from the sum total of environmental opportunities what 
it needs for its own specific internal environment, that is, the 
environment for the substance as such in all its subdivisions. 

The most obvious form of this selection is seen in ingestion. 
The cow eats plants, the tiger eats flesh, man eats both flesh 
and plants and a thousand things besides, the creation of his 
fancy; — but all eat as living substance seeking its own en- 
vironment, for these acts of the substance as organism are 
followed by kindred activities of the substance as such through- 
out its most minute extensions, the inter- and intra-alveolar 
filose activities being doubtless instrumental in attaining these 
ends, — and osmosis as a means is largely provided for by the 
physical form of the substance. (See above, New Structural 

[i2i] From a heterogeneous environment, common to all, 
each substance self takes, then, what will preserve, or create, for 
it its own special, internal, environmental conditions. Where 
the initial selection is made broadly, roughly, with seeming 
carelessness, or even wrongly by the substance as organism, 
the substance as such makes choice again within its safe pre- 
cincts, and again within its more secret haunts where we can 

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presently no longer follow it, extending and repeating the 
selection until all the protoplasm has chosen for itself, or 
passed by, or rejected, for itself, and secured, so far as the 
opportunities allowed, those special environmental conditions 
by which its general and its specific activities may be main- 

Later, when all the ingested matter has been passed upon 
thus, the substance as organism finally rejects and casts out 
all the leavings of the substance as such, together with such 
waste materials as have been thrown off by this during func- 
tion. In addition to direct methods of ejection, there are 
many indirect methods and also many special substance de^ 
vices for getting rid by more roundabout ways of deleterious, 
or cumbersome, or inutile, matter. Area after area of organ- 
ized physiological difference may be incited by presence of 
ingested food, to produce such secretions as will aid in pre- 
paring it for that diffusible state in which it can reach and 
be used as opportunities by the ultimate lamellar subdivisions 
of continuous substance in all parts of the organism. These 
areas are indirectly incited, not by actual contact with the food, 
and they act through their own existing conditions to aid in 
preparing what may later furnish renewal of their own powers. 

[122] Actively by means of its physiological powers, and 
passively by its chemical and physical nature and properties, 
the substance secures a double series of judgments on its 
internal environment. Then by new combinations and devices 
of areal differentiation, always on the basis of the foam struc- 
ture, it constantly extends its power of dealing as living sub- 
stance with the heterogeneous opportunities of environment. 

[123] To restate; in securing its own environment, the 
substance, both as such and as organism, exercises among 
the sum of opportunities offered it by external environment 
— a choice; and in this choice manifests its own character. 
The words " choice " and " selection " need carry no implica- 
tion of conscious action on the part of the selecting substance* 
They may be understood without further weight than is given 
the selection by a magnet of some steel filings from amongst 
glass and sawdust particles ; or a choice by the steel filings of 

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the magnet rather than of the fingers that hold it ; or even 
despotic chemical and physical interactions. A taking of one 
sort of thing from a number of associated and mingled oppor- 
tunities, — this is all the physicist or chemist biologist need 
suppose is claimed at this point. 

At times, the substance as organism selects or accepts 
materials which some specialized sentinel area within it re- 
fuses to pass, compelling an organism to reject them second- 
arily. There are certain areas which seem to be detailed to 
serve the organism thus and to pass for universal needs of 
the substance as such upon material admitted within it. In 
this connection the instance cited of expulsion of bacteria by 
a Vorticella, and of regurgitation by rotifers of their food, 
may again be brought into play. Every one knows the drastic 
action taken at times by difiPerent areas of the human body 
with respect to wrong sorts of internal environment. The 
most curious of substance devices to rid itself of undesirable 
internal conditions, is found in scavenger areas — leucocytes 
and phagocytes — which are so individualized that they select 
for their own specific environment what is deleterious to the 
substance in general. They are also used to take up surplus 
material of various sorts which interferes with normal activities 
of areas or the organism. Their function is doubtless homolo- 
gized for the substance as such by migrant currents and filose 
processes of the interalveolar stuff. 

That the organism makes even fatal mistakes at times in 
choice of its environment is true, but this is generally caused 
by lack of perfect subordination of the function of areas to 
the needs of the organism as such, for these by their very 
emphasis of irritable function may be led, for their own satis- 
faction, to wrong the rest of the mass ; or there may be in- 
escapable physical and chemical affinities which decide the 
matter, and the organism be taken by surprise ; or there may 
be an abnormal state in the purveyor area. Deleterious sub- 
stances that have gone too far to be returned are often hur- 
ried through the organism, or surrounded by some corrective 
or insulating substance which prevents their doing harm ; or 
again, if harm be inescapable, the powers of the substance are 

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seen in a new phase, for peculiarly suitable unusual secretions 
are poured forth as antidotal or cleansing agents. Local 
efiPorts of the substance to expel or escape or counteract inju- 
rious materials can cause disease or even death of the organ- 
ism, which then must be counted an indirect and not a direct 
result of the inimical agents. 

With regard to a taking up by living cells of useless or even 
harmful substances, especially during artificial experiment, cer- 
tain things are to be thought of. First, that the conditionings 
of the living substance must mentally be kept separate from 
itself. The foam structure, or local inclusions, may constrain 
areas into receiving such matter. And since alveolar cavities, 
even those of Biitschli's structure are used to hold waste, 
inutile, and excess, as well as harmful or unsuitable matter, 
entrance of the latter does not necessarily mean their selec- 
tion, or even their acceptance, by the living substance. It is 
to be urged also, that areas which for their guidance in taking 
up materials depend on certain specialised tactile senses, may 
be misled through these, whereas further within the cell or 
organism lie other local specialised centres whose selection 
may be guided by chemical reactions. What concerns us in 
all these cases is the subsequent action of the living continu- 
ous substance as such, or as organism, with respect to these 
intrusive things. // is therefore^ not alone physical reception of 
certain things by protoplasm in areas, which is to be understood 
as true substance selection, — but the full value of the most 
intimate processes, and especially the final verdict, of the living 
substance as such. 

That any nice adjustment of combined chemical and physi- 
cal conditions must be open to injury from chemical and 
physical violence — to consequent maladjustment which may 
become disintegration — is certain. That the living substance 
as knowable by us expresses an intricate, delicate, and most 
unstable set of molecular adjustments whose complexity is far 
beyond our analysis, can hardly be doubted. Is it not the most 
noteworthy thing that, in spite of this, it manifests so strong and 
persistent a hold upon its individuality ; that it can withstand 
such shocks and respond to such innumerable and kaleidoscopic 


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shif tings of conditions as often occur inside and outside it, with 
such plastic unity of phenomena. It is this mafvellous mainte- 
nance of its selfhood which seems to me more marvellous^ more 
worthy the powers of research workers^ than any abnormal states 
one can force upon the substance by virtue of its conditionings ; 
— especially as amongst the latter we are unable to separate the 
extrinsic from the intrinsic^ — or to say which reaction is due to 
these^ which to those. 

[124] Despotic as they seem from a familiar standpoint, the 
external contact relations of organisms and species are not as 
a rule the most important division of the sum total of such 
influences which g^ide its destiny. They are the conditions 
least under control, yet they are largely controlled by the 
substance's use of that other set of environmental conditions 
which it has gathered together for itself, and by which again 
it moulds new reactions for its intercourse, as organism as 
well as substance, with external environment. External en- 
vironment represents rather opportunities for the organized 
living substance. Internal environment represents at a given 
moment not only opportunities but intrinsic necessities for the 
substance. That the substance may be, to an important and 
often to a vital degree, influenced or even domineered over by 
external environment is indubitable, but to say this is not to 
say that the latter is its veritable control, or arbiter of its 
fate. As matters now stand, external environment dominates 
the living substance only in so far as it denies or inhibits it 
from using, or perpetuating, its own peculiar and necessary 
internal environment ; and the substance has devised many 
ways of eluding and defying external conditions in this. 

Stimulus from external environment is transitory, while often 
the substance reaction long outlives it, and that physical 
organization on which such reaction was effected may outlive 
both as it preceded both. In gross, the organized powers of 
the substance as organism are seen to be used chiefly, if not 
wholly, to obtain for the substance as such the material needed 
to continue or to produce, physiological function and physio- 
logical differentiation. And organization of this environment 
is secured by contact extension, and by repetition of the same 

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phenomena, i.e,^ the selection by the substance of its own 
specific environment. 

[125] This becomes plainer if we turn for a time to consid* 
eration of some ways by which the living substance evades, 
outwits, or defies external environment. In one class of such 
phenomena, the substance, finding external conditions adverse, 
simply sends them for the time being " to Coventry," isolating 
itself more or less completely from all immediate intercourse 
with them. In the lower forms, even in some so highly organ- 
ized as rotifers, this action is shown by the habit of encysting. 
In the higher forms, hibernation expresses the independence of 
the substance, and in both phenomena the substance shows that 
it is capable of withdrawing into itself where, by aid of its 
specific internal conditions, it can sustain the frown of circum- 
stances. Even the rhythmic nature of ingestion acts — which 
in different organisms cover widely different intervals — is here 
a most significant phenomenon, and the torpid state of the 
boa-constrictor (and of many savage peoples) after infrequent 
meals is but an exaggeration of substance habit which links 
these acts more clearly together with those whose object more 
obviously is to render the organism for variable periods inde- 
pendent of its external environment. In many cases and for 
variable periods, protoplasm itself is used to protect an organ- 
ism from external conditions, or to render it independent of 
these. The containing body purveys for and shelters, albeit 
often unconsciously or unwillingly, the contained body. This is 
the meaning of parasitism. The same thing is also expressed 
in most reproduction phenomena, notably in fertilization. Here 
again the thing of greatest importance to substance life and sub- 
stance habit of both host and guest in the dual life is their own 
internal conditions. In such artificial phenomena as grafts of 
areas and organs also, the substance seems often to live a truly 
parasitic life, continuing rather to select its environment in ac- 
cordance with its own accustomed habit, than to yield to the mass 
character of the host. Curious induced differences in habit 
of local deposit of reserve material, shown at times in regen- 
eration, may mean a use of inclusions prepared for diflFerent 
emergencies, or, abnormal secretive function in the substance. 

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[126] Thus much seems certain, that the power of the sub- 
stance to transport itself by contraction from place to place, 
either within or without the mass, either as organism or as sub- 
stance, either as a whole or as portions too minute for us to 
trace; its power by the same property to subdivide, or consoli- 
date, the alveolar inclusions of all the foam structures we can 
detect, and to transport them also hither and thither within its 
mass; and especially the power shown by the substance in its 
utmost simplicity or complexity of organization known to us, 
to select its own specific environmental conditions^: these 
things underlie at some point or other of their progress, all the 
manifold and complex phenomena seen in the animal world. 

All the phenomena of egg development seen in the Metazoa 
as all the phenomena of areal formation in the Protozoa, how- 
ever fleeting, may be expressed in terms of these results as due 
to organizing activities of the continuous element, tending 
always to more radical and sensitive selection of its environ- 
mental conditions as a basis for organized activities. 

The segregation of yolk and*pigment matter to certain areas 
during larval development, which is so finely shown in rotifer, 
starfish and echinus eggs as well as in many others; the con- 
verse segregation from many eggs with huge yolk masses, of 
the living substance, thus setting it free for its organizing 
development, are typical instances of such substance choice of 
physiological environment. The pursuit of light, heat, oxygen, 
or of other conditions among the opportunities of external 
environment show action of the organism as ^procuring device 
for the substance as such. The formation of contractile and 
irritable structures in response to stimulus, as described in a 
previous section, yields a still larger and more strongly typical, 
though closely allied, group of phenomena. Though due 
always to one final cause as stated above, protoplasmic organ- 
ization and reorganization has two aspects and is brought about 
in two ways. In one of these, greatest prominence seems to 
be given a placing of inclusions with respect to the substance, 

^ It is not claimed here that the substance can do this apart from its normal 
conditions or state of being, therefore experiments upon excised or mutilated por- 
tions of cells or areas have no bearing on the question. 

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and in the other it is a placing of the substance with relation 
to inclusions, as by migration, subdivision, or extension of the 
continuous element. 

[127] The formation of organized areas, of substance organs 
and of organism organs ; hibernation in ^1 its aspects ; motion 
from place to place in search of food or other environmental 
conditions which directly or indirectly affect the substance as 
such ; coalescence ; conjugation ; fertilization ; reproduction ; 
the birth of eggs or young ; the selection of proper external 
environment and the preparation of proper internal environ- 
ment for offspring substance by the parent substance, — as 
when an Ichneumon pierces a caterpillar, or a beetle rolls up 
balls of excreta, in which to place its eggs, or when a mammal 
feeds its offspring with milk : — all these seemingly very 
diverse phenomena are expressions of one and the same thing, 
— the selective power and habit of the substance as such and 
as organism with respect to its specific and necessary environ- 
mental conditions. 

Nor, as pointed out, do thes^ phenomena differ greatly from 
ordinary ingestion of food, or daily purveying efforts of the 
animal. In one case enough is taken for a seemingly secure 
rhythm of necessity, though a relatively short one, and in the 
others enough must be secured for a rhythm of privation which 
it is felt will be a long one. 

Among the lower forms, the creature does not seem to need 
to make special preparation but at any moment can enter upon 
such states of vital abstraction from external environment. 
When the organism realizes, or feels, adverse states of surround- 
ing conditions, it simply encysts or seals itself within a more or 
less impervious covering, and, suspending all action, waits for 
the proper moment to resume its life as substance machine. 
Or it throws its resources into the form of young which, from 
their physical form and physiological state, have far greater 
resistance to adverse conditions. Watching many of the Pro- 
tozoa encyst themselves, I have seen a relaxation of the normal 
alveolar arrangement to be common, so that physiological areas 
were mingled and the structure of Butschli became more 
regular, as in states preceding dissolution. . . 

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Nothing, I think, shows more clearly that the stable sub- 
stance as organism is merely a device for securing, and for 
elaborating and storing the specific environment for the sub- 
stance as such, and especially for the perpetuation areas of 
this, than the facts connected with the history of these latter. 
In the case of development of embryonic forms, we see the 
organism for long periods wholly independent of external envi- 
ronment as a source of supply, for it bears within itself all that 
is most essential to its growth, and without which it could not 
exist at all, no matter how favorable were all external conditions 
outside the eggshell, or membrane, or jelly-like covering it has 
been directly or indirectly provided with by the parent organism. 
And often the parent's own body wall shields and isolates it 
for long periods. 

[128] From the offered standpoint there vanishes that para- 
doxical mystery which surrounds preparation for larval or adult 
environmental conditions never experienced by an organism. 
Such phenomena express a reaction to already existing environ- 
ment. This is potent before opportunity to use an organ so 
formed arises, simply because it is in a sense independent of 
this. Such is the characteristic formation of cilia before the 
membrane of the egg is broken through; or of a double pelli- 
cular plate within the blastomere of an echinus egg before 
segmentation; akin also is the phenomenon of the beaver 
building his winter dam with fire-irons on a parlour carpet. It 
is all to be summed up in the saying that the substance's acts 
are based on its own internal conditions, and that all those 
structures which seem to precede function do not, after ally actually 
do so y for they are, in the first place, substance organs before they 
are organism organs. 

Peculiarly is the selection of certain portions of external 
environment by the organism for the substance as such, a 
similar manifestation of the same habit as a selection of cer- 
tain conditions by the substance as organism, either for itself, 
or for those portions of itself destined as newly incorporated 
organisms to outlive its older self. 

[129] The fact should be kept well in view that, before its 
separation, an egg or bud has been simply an organized area 

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of the parent substance, where were gradually segregated from 
the parental store some precious continuous substance with 
enough inclusion material of varied nature to serve the latter 
then and after for variably long periods of its career. Expe- 
rience of a disconnected set of facts, and a natural inclination 
to regard the state of being a unit or entity as of more impor- 
tance than the state of being continuous in kind, or character, 
with other units; have inclined us to think more often and 
more strongly of egg and sperm as future but imperfect organ- 
isms, than as what would seem to be their still more important 
character; namely, areas of differentiation of the parent organism. 
Like the lungs, heart, stomach, liver, — or even like the ovary 
and testis themselves, — they stand structurally for mass organs 
and for substance organs. And they contain characteristic 
quantities and distributions of the continuous substance in 
relation to specific materials. They should, I cannot but 
believe, be considered more from this point of view to be better 
understood. It will presently be spoken of how they arrogate 
to themselves the service of the whole machine and being 
served to the full extent of the powers of this, or to the satis- 
faction of their own intrinsic necessities as perpetuation areas, 
the machine may perish, if no better may be. As the daily, 
winter, or emergency store of nutriment is laid up for a periodic 
and rhythmically recurrent or intermittent set of conditions for 
the organism as a whole, so it is laid up in these areas for a set 
of conditions which are periodically or rhythmically recurrent 
in the race-history. 

[130] From this standpoint, the organism appears in the 
guise of a machine or device framed by the substance as such 
to secure its own specific internal environment ; especially to 
secure this for that phase in which it assumes the rdle of off- 
spring substance. There are frequently large and most com- 
plex areas set aside for the function of preparing and selecting 
from the sum of the parents' internal environment substance 
needed by the perpetuation or offspring areas. A machine 
within a machine is formed for use and service of the nursling 
substance. (See Parasitism, Fosterhood, Heredity.) 

The supply by parent organisms of specific internal environ- 

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ment or conditions for their offspring substance, and their 
labours to this end, seem still more wonderful when the 
ofifspring has undergone complete separation than when it is 
still an integral part of the parent. I should group the latter 
cases with those preparations made by the substance as embryo 
or larva for conditions not as yet unexperienced. 

[131] The attitude of the parent organism with respect to 
its offspring or perpetuation areas is evinced by its often 
directly renouncing its own specific environment or even its 
oVn continued existence for their supply. 

[132] A certain control of the living substance by external 
environment was the great theory set before us by Darwin. 
Yet it is not control by external environment of the substance 
as organism, but of internal environment by the substance as 
such which primarily rules the course of events and rules them 
most despotically, even to the point of independence and 
dissociation of the organism from external environment. 

[133] Substance habit, which in one aspect may everywhere 
and at all times be expressed by terms of organization of in- 
ternal environment with relation to the substance, has always 
been along lines of increased control, direct or indirect, of 
external environmental conditions; sometimes by increased 
differentiation, sometimes, as in certain cases of endoparasital 
existence, in the direction of greater simplicity of areal 

[134] In each act of the substance, we cannot escape the 
preformed machine, but at least in formation of a new machine, 
or new parts, or rearrangements, of the old, we see each step 
controlled by activities which seem to transcend the limit of 
power of the physical and chemical conditions of all visible 
machinery. All reactions of the organism to environment 
appear to me to express substance -reaction to pre-existing and 
specially prepared internal environment; to express the intrinsic 
powers of the substance under specific stimulus or direction 
from within rather than from without, — powers which, though 
they are for us finally inseparable from its physical and chemical 
composition, are still seen as separate from, and to great extent 
controlling, these. 

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Response to environment in character of its own peculiar 
intrinsic powers, — this is the power of the living substance; 
and, to repeat a point of radical importance, that response is in 
character rather than in kind. Sometimes these terms are 
indeed interchangeable, as when transmissory or sensory areas 
react to stimulus of activity, as contractility or irritability, of 
other areas of the living substance. To light, to heat, to 
impact, to chemical contact, the substance still makes response 
in character, over and above the immediate physical and 
chemical reactions which its physical form and chemical consti- 
tution necessitate. I have shown that we have at present no 
right to carry these conditionings so far as to make of this 
statement a petitio principii, 

[13 s] There is now for us a living, irritable, continuous 
substance of a protoplasmic foam; and there is its environment, 
internal and external; and these two sets of facts must at all 
times in our thought of them, as they are in fact, be kept within 
touch of each other, — never quite separable, yet wholly dis- 
tinct. It is the living substance in reaction to environment 
which has made and is still making the whole history of 
organic creation, — the substance's response to environmental 
conditions being always in character rather than in kind. 

[136] That standpoint which should regard animals or or- 
ganisms as disjunctive portions of an historically continuous 
substance — continuous in three dimensions of space, however, 
not in two; — which should use interaction to internal environ- 
ment to throw light on interaction of the same substance with 
external environment; which should in short read the phenomena 
of the organism to interpret phenomena of the substance, and 
use these to illuminate those; seems to me the most reasonable. 
The fact that our eyes look out rather than in, together with 
great defects of our optical tools, and the general opacity of 
living masses, have long withheld us from this point of view. 

[137] The substance both as organism, and as substance 
organ, secures so far as may be that which is needful for its 
energies down to its smallest pellicular subdivisions; and that 
substance as organism which has best cared for the substance 
as such, which has best borne neglect or privation inflicted 

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by external environment, which has been most plastic in its 
devices and activities, is that which is perpetuated. And its 
activities are developed by use along these lines. This presents 
a new aspect of natural selection. It is indeed upon the selec- 
tive power of the substance as such and as machine that the 
natural selection of Darwin must act. It must always be acces- 
sory after the act rather than an agent in the act, and even 
habit, individual or inherited, cannot be excepted. Through 
grasp of this basic fact of substance life, namely, selection of 
its own specific environment, individual, race and species habits 
are brought into a more unified and intelligible relation. 

The " struggle for existence " remains a struggle for food, 
using the word in its broadest sense. All struggles of the 
substance either as such or as organism for place and vantage 
in external environment as well as for the actual substances it 
holds, are to this end. Air, light, space, with more subtle con- 
ditions hardly yet understood in some cases, are also demanded 
by the organism because the substance as such requires these 
for its activities and internal arrangements. In the battle for 
these, such docile ingenuity and plastic contrivance are displayed 
by the substance, as more and more amaze research workers. 
The tendency of evolution seems to have followed lines of 
strengthening and conserving this aim and need of the Sub- 
stance, — to control its own environment. 

[138] Now food for the organism has commonly been taken 
to mean actual increment for it, either as new living substance 
or to replace portions of this which have suffered dissolution 
supposed to be a necessary result of characteristic activities. 
Certain remarkable correlations between the waste product, 
loss of mass or weight in the organism, the amount of energy 
displayed and the heat generating properties of the food taken 
in ; have formed the basis for a belief that food is used to repair 
waste of the actual living substance or to construct new 
quantities of this. The facts as to structure given here, make 
it quite possible and even plausible to suppose that we should 
not with certainty predicate this correlation between chemical 
and physical expenditure or supply and physiological activity, 
as meaning destruction or repair of the actual living substance. 

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We cannot predicate either the constitution or the final action 
of the living substance. What composes it, or how it grows, 
or is weakened, or dissolved, seems further from our knowledge 
than before, because all we have to base any theory on is 
chemical interactions between substances, none of which we 
can assert to be the constituents of the actual living substance. 
All the production of energy, all the waste correlated with it, 
may belong to the environmental conditions purely of the living 
substance. Its action upon them and even the waste products 
of this expenditure of energy may be wholly beyond our analysis. 
We may not yet have discovered the mode of analysis which 
can resolve for us these products, — if, even, they are chemical 
conditions known to us and within reach of our gross handling. 
It seems probable from these researches that out of the sum 
of all the innumerable chemical and physical conditions sought 
and appropriated by protoplasm, a few certain ones only are 
needed for actual perpetuation and reconstruction or construc- 
tion of the living particles, and that the bulk and majority of 
them are to serve as opportunities and stimulus for its internal 
powers and activities. It has been stated that owing to the 
physical form of the substance, growth of living masses can 
take place to an enormous degree without increase of amount 
of the actual living substance. Whether or no the living sub- 
stance itself is renewed by the myriad substances it accumulates 
within and about it, it is certain that these actually form its 
specific environment and the sum of the opportunities of its 
lamellar subdivisions, and that upon the selection and use of 
these both by the organism and by the minute portions of the 
protoplasm, depend the normality and organization of its activi- 
ties and the correlation of its phenomena, — more than this, 
the preservation of its life. In short, however we decide upon 
the question raised, the life history of the organism, from the 
time of its inception as an area of the parent mass, may be 
expressed in terms of supply for its internal environment. For 
the intrinsic powers and properties of the visible supposed living 
substance, are found to be alike wherever it was examined. Its 
specific areal modifications of function and products are to be 
taken apart from these. 

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[139] The whole series of structures which increase the 
range of organisms' intercourse with external environment, that 
is, the sensory organs, have peculiar reference to extension of 
the control of supplies for internal environment. The extension 
and intensification of irritable reaction, which are seen in the 
ectosarcal structures of sense organs, practically annihilate dis- 
tance and bring knowledge of supplies for which, if they must 
be in actual physical contact, the creature might wait long and 
perhaps hopelessly. As when a moth is drawn by odours too 
delicate for our perception, to its mate many yards away ; or 
a wild beast scents its prey, or an eagle marks his quarry, 
from afar. 

[140] According to the kind, and the distribution in envi- 
ronment, of its supplies, the character of the substance organs 
or structures of the organism are adjusted. In the plant and 
animal kingdom in their most familiar forms these things are 
strikingly marked. In both, the same general laws hold good, — 
that is the organisms select, not only what may serve for imme- 
diate use, but what may be laid up for use when the source of 
supplies is cut off from them, — or so lessened as to threaten 
the species by too fierce warfare amongst its units for the scant 
material : In both, the best efforts of the machine are spent 
for the needs of perpetuation areas: In both, the kind and 
quantity of the nutriment cause curious differences in the form 
and character, even the sex, of these areas. 

(a) Yet in each kingdom the characteristic choice is along 
the direction of those elements, or opportunities, surrounding 
it, the pursuit of which, as well as their chemical constitution 
when secured, tend to preserve and develope existing types of 
organisms. And with the character of substance habit in all 
these things goes the peculiar character of the ectosarcal 
structures, and the form of the machine as a substance 

(b) The animal kingdom has chosen the secondary, the un- 
certain, the motile, scattered, opportunities of environment. 
The plant kingdom, as a whole, has chosen to develope its 
powers along the lines of constant, stable, and, one may say, 
almost omnipresent conditions. 

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(c) Correlatively, the animal must develope along the lines of 
contractile and irritable structures, while the plant developes 
along lines of quiescent, assimilative, vegetative habit, and of 
chemical reaction. 

(d) To the animal, surface is of less importance than the quali- 
tative and quantitative structures developed within the mass. 
Economy of surface with concentration of powers and function 
will best serve it in the long run, in dealing with the evasive 
conditions on which its existence is staked. 

(e) In the plant kingdom the substance is extended and or- 
ganized in such a way as to secure for it as it rests passively 
in its place the largest possible quantity of light, air, mois- 
ture ; while its structures are so framed and grouped as to 
retain and control these supplies. In its reproductive phenom- 
ena, we see an approach to the more animal type of structure 
and function, because the more kinetic element must, like an 
animal, deal rather as contractile and irritable substance than 
as assimilative substance with environmental conditions. 

(f ) On the other hand in the animal kingdom we see often 
in the female reproductive phenomena a plant-like phase in 
which the assimilative and quiescent, or vegetative, state pre- 

(g) In plants there seems to be chiefly a quantitative exten- 
sion of function during growth and development. In animals 
while the powers advance along paths of both quantitative and 
qualitative emphasis during growth of the individual, the quali- 
tative emphasis is usually most notable. 

(h) In both kingdoms the full set of intrinsic powers of the 
substance seems to be alike and to be in great measure retained ; 
but their habitual expression differs in the two. Just as in the 
animal there are purely vegetative and assimilative areas, so in 
the plant there are markedly contractile, and in most, irritable 
states also, of the protoplasm. Indeed, in each cell area of both 
animal and plant these truths are characteristically repeated. 

(i) The extension of the animal's control over external con- 
ditions by means of its intensification areas will be discussed 
in full both as to means and manner, in my forthcoming 

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(j) In the plant kingdom^ the extra hoards of internal con- 
ditions become more rigidly adjusted to known rhythms of 
supply so that, as a rule, plant individuals are more readily and 
radically damaged by sudden environmental adversity than the 
animal units are apt to be, the habits of the latter being adjusted 
to meet intermittence in habitual sources of supply. 

It has been impossible because of rigidly limited space, to 
do more than indicate in this section the leading lines of argu- 
ment, and grouping of facts which have offered the standpoint 
discussed. Here too, as throughout the article, I have preferred 
to use as evidence so far as possible phenomena observed per- 
sonally, although there are innumerable established facts which 
might be cited, and the reader will, it is thought, find in his 
own mind stores of added confirmation. 

[141] To sum up in a few words the gist of what I have 
striven to make clear : The living substance down to its final 
visible subdivisions is seen everywhere seeking or selecting its 
own environment. All its guises and aspects, all its phases 
and phenomena are to this end ; and chiefly that the perpetua- 
tion areas shall be supplied with such environmental substances 
as will render them for long periods independent of the chances 
of external environment and then leave them prepared to resume 
the struggle for other similar areas in turn. 

[142] That so potent and radical a function of the substance 
should have areas of differentiation devoted to it, that it should 
have given rise to substance organs, or centres of such control, 
would seem a most natural thing. I believe that such organs 
were, indeed, among the earliest formed of all areas of differ- 
entiation, — excepting simple ectosarc. They are distributed 
through all living masses at short intervals, the first care of 
the substance in increasing its mass to any extent, or in in- 
creasing its scheme of areal differentiation, seeming to be to 
secure repetitions of just such centres of control or substance 
organs, and the surrounding substance in many cases, perhaps 
in most, seems to have lost the power to do without them, — 
just as a mammal has lost the power to do without its heart or 
lungs. I refer of course to the nuclei. Up to the present 
time there is ever-increasing evidence that they may rightly be 

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assumed to be selective organs for the substance, that is nutri- 
ment control areas. Whether these areas are this only or 
whether they serve also as substance ganglia, carry the form 
and function memory of the substance as such and stand for 
the nervous centre or brain of the cell area, must rest for 
future research, but the facts which connect them both with 
the cytoplasmic granules and with nervous structures have been 
pointed out earlier in the article. 


[143] I have said that many areas which as substance 
organs function markedly for organized contractility or irrita- 
bility, are formed of more or less unstably segregated proto- 
plasm in which the alveolar inclusions of the actively functioning 
substance are uniformly fluid. These inclusion fluids are 
carried from assimilating portions of the general substance, or 
received by dialysis, or even from wandering interalveolar 
foam. Such areas depend upon assimilative, areas for their 
special environment. To this extent they are parasitic. 
Whether they in their turn serve assimilative areas directly 
with their products, or indirectly by their activities, is aside 
from the immediate issue. Much interalveolar substance of 
Biitschli's structure forms a constant typical area of this sort. 
Parasitism is used here, in that broader, more genial, sense 
which does not preclude the idea of mutual benefits between 
dependent and supporting areas. In such organisms as 
amoeba where function and structure seem alike interchange- 
able, the substance forming true ectosarc may but a short time 
before have taken part in active assimilation areas, and even 
where ectosarc is a quite stable formation, its interalveolar 
foam may pass as an active carrier to and from these. By 
watching for hours the progress of assimilation in Protozoa, I 
have many times seen that there is at intervals a determination 
of fluid in considerable amounts to the food sacs — the tempo- 
rary stomachs ; and again a sudden redistribution at intervals, 
or a more or less steady draining, of fluids from such sacs into 
the surrounding protoplasm. These phenomena are mingled 

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with flux of Blitschli's structure and of interalveolar foam. 
Even in organisms having complex vascular systems, the work 
of these must be supplemented by just such minor or more 
primary dissemination processes, in order to reach areas whose 
continuous stuff has no immediate contact with vascular cur- 
rents. For much protoplasm in every organism, the continuous 
substance, as alveolar lamellae and as interalveolar channels, 
currents, and filose processes, both physically and physiologi- 
cally carries on this work, — which means to the substance as 
such just what the grosser vascular systems mean to the 

Every new formation of ectosarc however fleeting extends 
the parasitism of the substance as such. And every new 
area producing, or segregating, materials needed for contractile 
or irritable function of the substance elsewhere, acts as a 
device to furnish parasitic areas with their specific environ- 
ment, that is, with their intimate opportunities, or causative 
conditioning. Besides that physical restraint which the pres- 
ence of more solid inclusion matter may cause the contractile 
powers of the substance ; secretions, or chemical processes of 
primary digestion may also to some extent restrain, inhibit, 
or intermit these. They may be necessarily alternative or 
secondary results. The converse may also be true. With 
regard to Biitschli's structure, ectosarcal areas stand often for 
secretory habits of more or less pronounced character, for in 
this structure, as stated before, all sorts of materials are 
deposited, and an organized number of vesicles having kindred 
deposits would mean, or indicate, a local habit of the general 
substance as such. If fluid, such deposits form physical 
opportunity for organized contractile and irritable function. 
In other cases these inclusions may serve to inhibit character- 
istic activities of the substance, and then this will commonly 
be withdrawn by degrees to more free conditions. In many 
cases parasitic areas take from the general store, or from 
special areas, matter which is useless or positively hurtful to 
the substance, or which is to be transported elsewhere, or held 
in constraint for coming substance organs. (See Heredity.) 
Such are many wandering cells of Metazoan systems. These 

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have a curious individuality of their own, — they are like 
organisms within the mass and act as carriers with such seem- 
ing sentience, with so surprising a sympathy of response to a 
host of internal needs and states of the general substance 
that it is only when one looks at them as free connective tissue 
areas that one can at all understand their relations to and with 
the general substance. Such service as they yield in grosser 
expression is supplied the substance as such more radically by 
migrant currents, by filose processes, and perhaps by chemical 
or physical conditions. I have watched in situ in soft shelled 
crabs, certain of the leucocytes take the form of bi-fiagellate 
monads with variably protoplastic posterior region, and travel 
about thus, constantly insinuating with some appearance of 
force the tip of their long flagellae which were sharply pointed, 
into the tissues about them. Whether they brought sub- 
stances or took them away or stimulated excretion from these 
tissues could not be seen. One would perhaps think them 
parasitic forms were it not for their frequent metamorphoses. 
In blood drawn from the crab they quickly change to the 
Heliozoan like state and aid in the coagulation phenomena. 

[144] Amongst organisms examined, from Protoplasta to 
highly organized Metazoa, the complexity and number of areas 
laid down for organized contractility and irritability appeared 
to be correlated with number and complexity of devices for 
assimilating, and for transforming assimilation products into 
most varied materials, all of which when presented to the 
substance as such were in dialysible form. (The male rotifer 
and similar cases, as of spermatozoa, do not form a true 
exception here, — see following sections.) Looked at from this 
standpoint^ cell division^ and, more broadly yet, areal differen- 
tiation appears as chiefly a multiplication of ectosarcal areas or 
organs and, with this, of a nuclear machinery to control their 
specific supplies. 

Parasitism of the substance being an established substance 
habit, it like all other such appears throughout the manifold 
relations of the substance as organism. And here, as in 
other substance habits, there is in higher organisms progressive 
increase and extension of it in all the relations of the sub- 

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Stance. Such parasitism as does not belong with phenomena 
of true fosterhood may arise in varied ways. It may occur 
either as direct selection of another animal as a chosen source 
of supply, or accidentally from ingestion of one organism by 
another. The organism so ingested may find its new sur- 
roundings so much to its purposes, that is, so much in accord 
with established substance habits and organs of its own, that, 
far from succumbing and becoming chemically disintegrated, it 
is as it were enclosed in a factory or source of artificial supply. 
Instead of serving as food for its captor, it uses the latter 
either as endosarc, or purveyor organ, or actually and directly 
as living food — devouring its tissues. Or, it may pass to 
some special area and there merely filch from the prepared or 
selected food what its own uses require. It is not strange 
that under such circumstances the plastic substance should lay 
aside or modify substance organs for securing and preparing 
its food, while retaining and accentuating those which best fit 
in with its new set of opportunities and compulsions. In a 
retention of their own individuality by grafts or transplanted 
structures, one sees that part of an organism may continue 
to maintain its selfhood while fed by another organism, of 
which by substance parasitism it becomes physically an inte- 
gral part. // shows again the importance of the substance as 
such^ and that the local form is not merely part of a whole 
organism as such^ but primarily and radically an expression of 
substance habit, (See Heredity.) Such facts throw light also 
upon the physiological differentiation of substance organs with 
continuity of a physical and physiological nature between 

[145] Once established as dependent, the parasitic sub- 
stance, whatever its habit, forces the supporting substance or 
organism to play the part of foster parent and itself resumes a 
r61e which as fosterling substance, or perpetuation area, it 
once played. For among all the multiplex dependent areas, 
those most unmitigatedly parasitic and exacting are the per- 
petuation or offspring areas. By a parasitism which is in 
harmony with its fundamental structures and habit, or which 
releases it temporarily from great enforced strain of its condi- 

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tionings, or which enables it to make fuller use of these, the 
substance as such and as organism seems to profit in renewed 
vitality. These conditions are to be found very commonly 
alternating with converse states of initiative activity of both 
the substance and its gross expression in organisms. The 
more vicarious means of supply can be extended, the more the 
substance may expand and expend its own peculiar powers in 
self-expression. But unless states in which an endosarcal set 
of habits are peculiarly fostered alternate with ectosarcal self- 
expression the organism or area becomes by so much degener- 
ate ; and if the latter outrun the former it becomes exhausted. 
The value of such alternation is probably at the root of that 
flux of living substance in organisms which has been so much 
emphasized throughout this paper. Here again one sees sub- 
stance habit and its conditionings apply to organs and to 
organisms — peculiarly in reproduction phenomena. 


[146] The word parasitic having by a certain frequent use 
associations which in some connections are jarring, it should 
suffer as idea a transformation of verbal form gracious enough 
to follow that of the fact into the beautiful phenomena of 
parenthood. Dependent perpetuation areas may be called 
fosterling areas ; the supporting substance foster substance, 
area, organ, or organism. Of all areas differentiated to live at 
expense of other parts of an organism, there are none so 
grossly egotistic, none which so take all and keep all for them- 
selves as the perpetuation or fosterling areas. From their 
inception, for variable periods, often covering the whole term 
of their existence, they receive largely from the foster sub- 
stance in many of its phases. It is common for a parent 
organism to support a special machinery whose work is alone 
for the offspring area. The whole parent itself as organism 
may be regarded as serving as a substance device for securing 
and preparing the materials needed by the perpetuation sub- 
stance. For the pampered fosterling areas the rest of the 
substance may go more or less lean and hungry. To them 

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may be sacrificed not only the general mother substance but 
the father organism also ; more than this, many potential 
fathers may perish in warfare merely that one such area shall 
have the best of the material it needs. There are even cases 
in which the entire father organism is but a servant area for 
the fosterling substance, having existence, and possessing 
no other parts and powers, than are necessary for this end 
only. It is but the sham of an individual, neither eating nor 
seeking food for itself. It acts only to secure for the perpetu- 
ation substance it carries in the form of sperm, certain specific 
environment this needs. It is a messenger foster area of the 
mother organism. All its structures are adapted to this ser- 
vice and it is an emphasized expression of the true meaning of 
perpetuation machinery and areas. Such are male rotifers, — 
they pass through an egg phase but remain free swimming 
foster areas of the mother organism. They never advance 
beyond a true parasitic state, — they live and move wholly by 
the specific environment the mother machine stored up within 
them, and they perish shortly after accomplishing* their mission, 
that is when the sperm has been deposited within the body 
cavity of the mother through a hole the tale spike of the male 
pierces through the body wall usually about the shoulder region. 
They possess no means of receiving or assimilating food, are 
chiefly if not wholly ectosarcal, and are largely motile, tactile, 
and perceptive areas. They have, let me repeat, no true self 
expression as individuals, not even the semblance of it common 
to organisms ; they function wholly as separate areas of the 
mother rotifer — and for the race. They seem to me from a 
hundred facets to illuminate the place of the mdividual in the 
history of the substance. They form also a fine illustration of 
the extent to which what one might term vorsichtigkeit of the 
substance extends. A whole generation, speaking from the 
usual standpoint, is interjected by way of providing that next 
the mother generation with all its material, for the sperm go to 
perfect a generation which will be truly collateral with that 
formed by the males themselves. This would greatly disturb a 
genealogical record, for the child of one parent seems to be the 
grandchild of the other. In such a case the male is doubly 

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parasitic upon the mother generation, — as a free perpetuation 
machine, and for the sperm (which does not tndy belong to 
the male) and for specific environment for its future develop- 
ment and life. As stated in a preceding section, the male 
element is largely dependent always upon the female for its 
specific environment. Other points of extreme interest in con- 
nection with the male rotifers are these. They are formed of 
eggs about one sixth only the bulk of ordinary parthenogenetic 
eggs producing females ; they are produced about six times as 
fast ; they are formed at times when the supply of food is 
threatened by climatic changes and when the parent organisms 
need all their reserve stores for formation of the huge winter 
egg which is fully six times the bulk of the usual summer egg. 
The male does not even diminish the possible food supply for 
the parents since it does not, so far as known, ingest. All 
these facts seem to me to make clearer the true relation of 
organisms to the race history of the substance, and of ecto- 
sarcal areas to substance habit. The facts were personally 
observed in a number of forms but most continuously and 
repeatedly studied for a number of years in Megalotrocha alba 
and M, sp. ig. 

[147] An enormous proportion of the habits and instincts 
of the whole living kingdom exist in relation to fosterhood 
phenomena, especially with regard to the perpetuation areas. 
Among the lower forms are many which retain for variably 
long periods, or always, vital protoplasmic connection with their 
fosterling areas even after these are manifestly self supporting. 
Now just as, by qualitative extension or intensification of the 
irritabflity of the living substance, more primary organs of g^oss 
and direct contact found in lower forms are transformed among 
the higher animals into space-bridging organs of sense percep- 
tion ; so here again the same phenomenon of transmutation of 
structure and function is repeated. Long after actual space 
and time separation the parent can thus extend recognition and 
assistance to its fosterling substance and still continue to subsi- 
dize its own supplies or functions. 

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[148] ^ Up to this point it has been cumulatively shown that 
cell phenomena are underlain by such phenomena of the con- 
tinuous substance as would seem to inhibit us from using cells, 
even broadly, as primary units of physiological organization ; — 
the new facts urging us to trace substance phenomena in a 
physical and physiological continuity throughout all parts of 
organisms ; to ignore cell limits, except as they fall within this 
interpretation ; to see in cell walls and in nuclei local and even 
temporary substance organs belonging primarily to the mass 
and but secondarily to cells, their curious repetition being taken 
in relation to general needs of the substance as such rather 
than as parts of cells as units of structure ; — in short to study 
cells as localities in a mass organization of the continuous sub- 
stance and as local expressions of substance habit in a signifi- 
cantly common grouping. From this standpoint and by use of 
the naYve method of thought approach, the grosser organs and 
structures of organisms become replaced by an intricate maze 
or web of substance organs. Organs no longer appear as 
compounds of certain different sorts of cells, but as a complex 
of minute substance organs whose multiplication baffles even 
the imagination, for they not only extend in a lessening series 
into the invisible subdivisions of the continuous substance but 
are constantly being transmuted into new structures. 

The physical form of the contractile irritable substance 
offers peculiar opportunities for physical organization, for each 
vesicle is a nucleus of varied causative power and may form in 
the series of emulsive foams a radiating centre of influence 
along several, or along many, lines of continuous substance 
and of vesicles. Each local aggregate of inclusion fluids of 
kindred sorts forms at once a true substance organ whose pos- 
sible structural and functional values are many. The physical 
form of the living substance of itself may be said to initiate or 
even compel a formation of substance organs, the chemical 
nature of the constituents to extend aiid qualify this natural 
compulsion, while activities of the living matter may expand, 

^ Whole section to be read under this number. 

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modify and restrain, or with aid of the mechanical conditions 
may freely form other such organs. The minutest of sub- 
stance organs, whatever their origin, express potent specific 
conditionings for living lamellar films and foams. Other areas 
directly or indirectly profiting in any manner by them, they 
acquire the value of organs of the general substance ; the mass 
becomes functionally, as it is structurally, enriched ; the or- 
ganism becomes, though perhaps but fleetingly, by so much 
more an organism. In more stable retention, in maintenance 
or iteration, in multiplication and in varied extension of sub- 
stance organs, the structures and organs of organisms as 
familiarly known to us have their birth. These latter group- 
ings come secondarily, incidentally as it were, to form and to 
belong to what is commonly known as the organism. They 
were and are primarily for the substance as such. I believe it 
may be said of substance organs ; they are, — therefore the 
grosser organs, — therefore the organism. Through unity and 
harmony of form and interrelation of function in substance 
organs, and through secondarily acquired interdependence of 
their compounds, there arises a most specious seeming of prime 
end or motive in these latter, and especially in the organism 
they form. (See Habit.) 

But besides being structural complexes, organisms are, even 
more, functional complexes. From the same standpoint and 
using the same method of approach, an organisih expresses 
itself functionally as a maze of sequent phases, composed at a 
given moment of many lines of phenomena which are in 
closely interwoven extension, and even anastomosis, in three 
dimensions of space. The life history of cells and again that 
of organisms and still again that of the race, repeat in grosser 
terms the same truth. Of minute groupings of function in 
substance organs are built up all the functions of cells, of 
organs and of organisms. But the complexity of these outruns 
structure many times, for a given vesicular structure may 
be the seat and cause of, or opportunity for, many diverse 
sorts of function. Physical conditions due to mode of lamellar 
distribution can alone give rise to one set of phenomena, 
chemical interactions may be manifold and either or both of 

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these sets of phenomena may be locally isolated or have im- 
portant relations with other substance organs. These physical 
and chemical activities may alternate with, or be controlled or 
pretermitted by, physiological activities which may also make 
large use of them as opportunities. Of causative interactions 
amongst any one alone of these sorts of phenomena, volumes 
might be written, and still incalculably much remain for coming 
centuries of research workers to write ; — and all three sorts 
are actually interwoven and mutually transmutable as cause 
and effect. 

Constant watching of function in the living substance gave 
the following results. The function of any substance organ 
would seem not to be due to peculiar powers or properties of a 
locally segregated portion of an organism's substance ; but to 
exist as specific temporary relations of a universally similar 
substance with its specific conditionings. All the facts gath- 
ered as to structure and habit of the living substance seem to 
me to indicate that it is indeed identical in powers at all points 
in organisms ; such seeming differences as it shows expressing 
for the general substance, local habit of selection, organization 
and use of chemical opportunities or inclusion material in rela- 
tion to itself; and for the local substance, specific chemical, 
physical, and physiological relations with these, influenced 
more or less by various controls due to function of other areas. 
Endosarc becomes ectosarc and there functions in character 
of that mode of organization and opportunities. Ectosarc re- 
turns to endosarcal areas to function there again in character 
of the latter. These things are true of the substance com- 
posing higher organisms too, for even here the substance was 
seen to exchange special organized states or conditions for 
those more primitive ; and, correlatively, its functions for more 
primal habits of activity. This truth is already partially 
known in a grosser form of its expression as a " conversion of 
function ** in organs ; but in the basis of these, in the true 
substance organs, it is almost, perhaps quite, universal. I have 
shown that in minuter structures the substance is in a state of 
local flux, and that stability of visible structures is but a mask 
for mutability and mutations of both function and finer struc* 

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tures ; also that such stable dififerences as are had by staining 
reactions are attributable, in the continuous substance as well as 
in Biitschli's structure, to differences of inclusions and of phys- 
ical states of the lamellar substance, rather than to chemical 
differences in the true protoplasm. The continuous substance 
of contractile or irritable organization, functions often in secre- 
tory or excretory manner as well, and in addition to its organ- 
ized modes of expression, frequently intermits these for, or 
mingles them with, primitive protoplastic manifestations. The 
migrant protoplastic substance can maintain the finest visible 
substance organs while yet retaining diverse other character- 
istic modes of function. Organs, as we have known them, are 
expressions of the living substance in some particular way, or 
ways, but with enormous reservations of power and possibility. 
Structure as hitherto known, seen thus, takes on the aspect 
of locally maintained deposits of specific inclusion matter. 
Function expresses certain local or general habits of the continu- 
ous substance with respect to or in relation with these mate- 
rials. Manifestly, in, and by, no known structure, is the 
substance expressing at a given moment its full powers, or 
asserting a prime local difference in itself. The utmost expressed 
thus, is mass habit of local deposit and use of certain materials, 
and within this expression is still a wide range of structural 
and functional difference and variation, not only in individuals 
of one family, but in each unit from moment to moment. 

Another thing of note is that the living continuous sub- 
stance organism ignores, transcends, controls, and rules, as 
well as directly responds to, its internal opportunities in any 
visible structure just as does the organism as such in relation 
to its external opportunities. The continuous substance fol- 
lows internal controls and promptings not to be associated 
wholly with, or wholly accounted for by, any structure we can 
trace ; it functions according to suggestion or control of more 
intimate conditionings, — yet always in such a manner as to 
repeat in its true activities the phenomena seen to be asso- 
ciated in visible structures with vesicular modes of organization 
of the elements. These intimate conditionings are variously 
derived and may give rise to yet others. 

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Most important of all does it seem, that the function of all 
substance organs, simple and compound, is more or less inter- 
mittent, evanescent or recurrent, the recurrence having com- 
monly a rhythm which may be for the local organ statedly 
self-supporting, statedly dependent, or uncertain because con- 
tingent on uncertain aid from within or without. Separating 
out from the continuous network of phases, those lines or 
rhythmic modulations formed by linkedly recurrent phe- 
nomena, they too weave in substance life history a coarser web 
or maze of sequences and inter-relations. Sequence lines of 
minute substance phenomena form part of larger rhythms, 
these in their turn create still broader organic inter-relations. 
Each phenomenon of the local substance organ is not in each 
for itself alone but for all relations with other parts of the 
vesicular organism. Again, each phenomenon is not for imme- 
diate time alone but may be vitally, or indirectly, causative or 
restraining for remote as well as for directly sequent phe- 
nomena. Often numbers of other linked or associated phe- 
nomena must arise and pass before its fulfilment is ripe. The 
end of any substance organ and its function may lie in the 
developmental history of a coming generation, or even of sev- 
eral generations in advance, for the linked causal nature of 
substance phenomena is very far reaching. Direct results of 
physiological activity may have a purely physical or chemical 
value, or both, and from these again may be built up new 
physiological manifestations. Whatever its origin, — to what 
end a substance organ of minute extent and simplest vesicular 
organization is to function, or by, or through, what remote sub- 
stance activity it has been prompted, must at all times be in 
full beyond the grasp of the wisest man. No matter where 
situated, nor within the limits of what broadly marked organ 
of the organism it may lie, it may have, or be, a remote result 
or cause, having little actually to do with its immediate func- 
tional surroundings. The instability of substance organization 
and the migrant protoplastic nature and habit of the substance 
alone make clear that results or products of local structure and 
function cannot be limited in sequent efifects to the same 
locality. There is also much distribution of local products of 

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organization throughout all living masses. It is certain that 
the local functions of the substance belong not alone to the 
locality, not alone to the cell, not alone to the tissue or organ, 
— but to the mass; and again that they belong not alone to 
this, or even to its direct descendants, or to the family, but to 
the race. It is the life history alone of the substance as race 
substance that can reveal the full significance of even fleeting 
local organizations. 

With all its complex interrelations which seem to form so 
complete a cycle, the individual is in truth but a fragmentary 
part of a vast complex modulation of structure and function. 
However separate as mass from its race relatives, it is abso- 
lutely inseparable as part of the web of sequence in race phe- 
nomena. The strange incomplete, yet redundant, complexity 
of sequence we call an organism, is, as pointed out earlier, but 
a ceaseless becoming, of which the adult, or alL stages, form a 
recurrent grouping of phenomena — a kind of rhythmic wave 
in continuous habit. It is thoroughly transitional, for in it 
nothing truly begins, and nothing truly ends. But the being 
is more obvious than the becoming ; the structures and phases 
when viewed in gross appear more stable than evanescent ; — 
and so the gross result produced upon our perceptions is a kind 
of typical stability or homogeneity. What is seen and described 
at any given time is, however, a net result of composite impres- 
sions beyond our analysis. Form and structure are as much 
illusions bom of our o^yn limitations of perception as has been 
our conventional mode of seeing and representing apparent 
modes of motion. Thus have arisen our types, by convention, 
and yet in a true way after all, if thoroughly understood to be 
symbolic. The form of flame, or wave, or fountain can be char- 
acteristically expressed spite of all the local evanescence of 
forming substance. But this is form symbol, not form fact. 
Just as in Protozoa a characteristic form or forms, with general 
relations of parts, is maintained in the mass notwithstanding 
ceaseless flux of formative substance, so is it in all organisms, 
only, in most, the structures and phenomena which control our 
attention and the grosser interdependences of these are less 
patently unstable. And it must be remembered that with the 

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same powers required to see the flux of Protozoan forms, the 
flux of substance in Metazoan forms can also, with greater 
patience, be observed. Within and through the most complex 
mechanism of cells, the continuous substance or formative 
agent moves restlessly to and fro expressing itself as living 
foam in countless diflFerent ways. Primarily protoplastic, it 
builds up in relative localities of living masses specific deposits 
of non-living material and maintains these in a general way, 
without for the most part hampering its total liberty, merely 
using them to make itself more free in and for self-expression. 
In developmental phenomena, one sees the machinery of the 
embryo formed by successive progressive shufflings of inclu- 
sions already placed in it, and by more and more searching 
rearrangements of these in relation to delicate pellicular mem- 
branes of its living continuous substance. As differentiation 
advances, area after area becomes markedly possessed, though 
more or less unstably, of its own specific inclusions and certain 
substances become wholly restricted for variable times to cer- 
tain cell areas or to minuter areas within these. Nuclear 
division phenomena seem to me to stand for subdivision of 
specific conditioning material, for repetition of a local control 
machinery having to do with general needs of the substance. 
Whether as a ganglion or as nutritive control it is not possible 
as yet to say certainly. Probably as both. 

The life history of perpetuation areas, their dependent rela- 
tion to the foster machine or organism ; their relative im- 
portance in time of famine as compared with other areas all 
clamorous for food ; their long periods of reaction to the 
internal environment provided for them ; their relative vigour 
and vital power as compared with the parent organism when 
first completed for self-support, or before this even ; — all 
these things seem to me to declare that in these areas the 
living substance is kept young or re-invigourated by sparing it 
for a time all wasting activities and by feeding and cherishing 
it in all ways, even if to do this be to sacrifice the parent 
machine. The business of the substance isy after all^ self-con- 
tinuation, rather than continuation or reproduction of an existing 
grosser organism^ which is but incidental to its prime habits. 

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The formation and storing of perpetuation areas represents no 
break or radical difference in the general habit of the mass 
substance. It is to be remembered that food secured by the 
organism is always for activities of the future, for even remote 
contingencies |is well as for immediate or certain rhythmic 
needs. That some of these needs will not arise till a portion 
of the existing mass shall be leading an independent existence 
matters little, for the sequence of structures and especially of 
function has still for this a certain and a true continuity. 
Considerable interval must always elapse after taking food 
before the general and local substance has use of it. A stor- 
ing of perpetuation areas with specific environmental condi- 
tions needed for its sequent phases and phenomena is, 
therefore, one with all the phenomena of distribution of 
materials throughout the mass, one with all preparation made 
by a larval stage for other larval, or for adult stages. And 
indeed the *' adult " stage in many cases becomes absolutely 
parasitic upon the ''larval" stage. In such cases it is the 
whole and not merely a portion of the mass that becomes 
fosterling, for which the precedent phase acts as foster parent. 
Some adult phases are as absolutely dependent on this as the 
ovum is upon the mother organism. In pupa states again, the 
whole organism passes through an egg-like phase, becoming 
regenerate and transformed. The male rotifer must again be 
cited in this connection as an illuminating form. By regard- 
ing the organism as a recurrent grouping of substance organs 
and substance f unctionings, — as a rhythmic set of substance 
habits, — the whole living universe comes more within one's 
comprehension, because expressed in more unifying terms. 

But it is manifest that while the creature we know, is built 
up of substance organs, it can hardly be taken to be by itself 
the whole meaning, a mere summation of those found in it. 
In each organism there are probably vast numbers of true 
substance organs existing and functioning with little or no 
reference to it, except as they are indirectly dependent upon 
it as general purveyor. Each of these may represent some 
attained end or cast-off function which will disappear — van- 
ishing characters in process of elimination from the race 

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history ; some initiative characters one day to mark a new 
species — incidental but potent beginnings ; some dying 
results of exhausted or unused opportunities ; some products 
of an overloaded, overstimulated state which will suffer read- 
justment ; — any of these things each may be. Whether an 
evanescent possibility, or initiative of some new series of fruit- 
ful structural deviations, whether to be used by the substance 
as a physical lever, a chemical cause, or perhaps as a mere 
storehouse of opportunities, or prison for adverse or detri- 
mental influences ; — such is in each case, and in each moment 
with respect to the same vesicular structure, our problem. 
One may be watching the last falling leaf of some great tree 
of biological sequence, or the forming of embryonic leaf and 
root of another such. The organism would seem to be more 
than we have thought it and less than we have taken it to be. 
It is an incident rather than an end, but as incident far more 
marvellous and comprehensible than as prime end. A most 
complex and compound race organ, it represents a recurrent 
grouping of substance habits as to local deposit of material 
and local relations with these. Of such are formed all like- 
nesses of character between organisms and their grosser 
structures ; likewise all differences whether of individual, 
species, genus, family or race. But within the limits of the 
most closely resembling descendants there is room for enor- 
mous variation, as well as of mere diversity of substance 
organs, both as to their extension and complexity, without pos- 
sibility of our detection as yet. Possibly in many cases with- 
out any essential difference resulting. This is, it seems to me, 
a most natural result of the mode of organization and growth. 
Of such of these variations as best fall in with the opportuni- 
ties of the organism as race organ and race substance, of such 
as least interfere with, are most closely allied to, or most in 
harmony with, established tendencies of race rh)rthms as well 
as those of the individual, with latent as well as with patent 
lines of substance phenomena ; grosser variations will probably 
be built up. For thus, it seems to me, natural selection must 
act. As substance organs arising from mere physical incident 
can become true mass organs, so the organism arising from 

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actions and interactions incidental to aggregate substance 
organs, can become a race organ. Considering the strange 
unlikeness between different phases of the same unit, and that 
of sexes to each other ; the marvellous metamorphoses also 
common ; the alternation of generations ; the single, dual, 
triplex, or multiplex, relation of parent and offspring organisms 
with their common and enormous differences ; the colonial and 
social groupings of diverse creatures ; — the place of the unit 
becomes, I think, unmistakably that of a certain organ of the 
race substance, incidental to the race history of that substance. 
Such structural and organic basis as has been used for group- 
ing animals, now seems gross indeed when brought into con- 
tact with true substance organs and structures. The difficulty 
of understanding how organs which could not be of use to 
the organism until already far advanced could be built up by 
natural selection, is thus lessened, for they are to be thought of 
first as substance organs and then as organism organs. And 
their function in these two capacities may often be widely 
different. We may know, it may be patent, how any group of 
substance organs serves the organism as such, or acts in a 
group of interrelations between its parts ; but we cannot be 
therefore sure that we understand what part this organ plays in 
the life history of the substance, especially as race substance. 
Likeness of the adult offspring to the immediate parent was 
so common a phenomenon in most accessible groups of beings 
that it assumed at once in man's mind an undue importance 
which it has ever since by tradition retained. It is rather 
incidental than of prime importance except as a general 
expression of certain broad recurrent groupings of substance 
habit to form a race organ. A useful fact — and one that has in 
general looked like a final fact when the history of units was 
regarded — it has seemed indeed to be an end to which all the 
wealth of power and adjustment known in the organism was 
straining. To say that it is a natural and necessary result of 
the fact that the same powers which formed the parent are in 
operation at its birth, would be, I think, an inadequate state- 
ment, for it may be a result of powers and processes which 
were wholly or largely in abeyance when the parent was 

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formed, and the powers and processes which formed both it 
and the parent are by no means to be limited or defined by 
those results, nor have they thus wholly expressed themselves. 
The individual may rather be said to be one recurrent result 
of conditions that guide substance habits along lines produc- 
ing race history. The rhythm of that recurrence may in all 
or a variable number of its parts include many generations. 
Atavism as commonly known is but a grosser expression of 
this truth about substance habit. The father element con- 
tributes his portion of the web of phenomena of substance 
habit and substance opportunity ; the mother hers. Both 
doubtless contribute some of each sort of conditions as well 
as of living substance, but the mother, as a rule, patently sup- 
plies most of the immediate inclusion opportunities. Sex is 
determined, it seems to me, at some variable time by existing 
internal opportunities which decide the sort of race organ 
most possible, or most in harmony with these. The race organ 
has other meaning than immediate self-expression of the sub- 
stance, but has also its relations to existing external oppor- 
tunities as well as internal and these become known to and 
recognized by the sentient mass substance. 

From this standpoint one no longer thinks of " transmission " 
but of transitions; one no longer wrestles as Jacob with the 
angel, with the problem of transplantation and distribution of 
representative and determinant biophores, — but one sees 
spread out before one, sequence of phases and phenomena, 
origin and maintenance of substance organs and substance 
habit, and the maze of intermittent or recurrent rhythms these 
form; — one sees the whole race history of the substance as a 
vast, web-like, compound grouping of physiological as well as 
physical and chemical phenomena, in three dimensions of space. 
The individual becomes a single chord in a great orchestral sym- 
phony. At this standpoint one finds oneself suddenly relieved 
of the burdeh of mental necessity or yearning for any theory 
of heredity, for such "heredity" appears here to be but a 
phantom difficulty, a result of artificial separation of portions of 
the web of sequences, a term forced upon us by an inherited 
standpoint from whose necessary limitations we have suffered 

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If the difficulty does not truly disappear as it seems to do, at 
least it no longer stands between one and simple organized 
effort, it no longer thrusts itself between one and the subject. 
Here, it seems less urgently necessary to speculate of the how 
of heredity, than to watch the living substance and learn its 
varied sequences of self-expression. 


[149] To substance habit, then, must we look for all likenesses 
and differences found in living organisms. This might seem 
merely an exchange of terms, leaving the root difficulty un- 
touched, were it not that our new facts make it possible to 
separate habit into more fundamental sets of conditions and 
functions; and this is useful, for, in the substance as such as 
well as in the organism, habit is a most complex state of 
things.^ In terms of my results habit of the living substance 
in all its relations may be described thus: 

Habit expresses certain characteristic vesicular groupings of 
the living and non-living elements of protoplasm; it expresses 
also any or all iterated or characteristic primary, secondary, or 
remotely linked, interactions of these conditions amongst them- 
selves, or with any existing opportunities, compulsions, or 
stimuli within or without the mass or area ; it can also mean 
simply a state of actual or cumulative preparation, with intrin- 
sic powers, for such interaction whenever it shall become 

Protoplasm in which no structure beyond a nucleus is seen, 
is commonly called " undifferentiated " or " the simplest form 
of organism." All visible structures and their associated 
functions, all visible activities of mass or area, we are used 
mentally to bestow upon or refer to the " organism." When, 
for instance, the cuticular alveolar layer of a Vorticella 
responds with organized contraction to local stimulus, we say 
'the Vorticella contracts,' just as when a dog's Ifeg moves we 
say * the dog moves his leg.' Because cuticular contraction 
often removes the Vorticella from harm's way it has seemed 

1 Organism will now be used in a limited sense to imply only those parts and 
functional interrelations heretofore familiar to us under that name. 

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a purposive habit controlled by the organism. By changing 
this standpoint, — by taking structure and functions of vesicular 
organs in their relation to the substance as such and to each 
other as organs of this, — much will be gained. In any visi- 
ble structure the continuous substance is a protoplasmic foam. 
From or through this must the coarser vesicular inclusions be 
built up and maintained, either chemically in situ by the local 
substance, or as deposited products of the general substance, 
brought, it may be, from some distance by physical or physio- 
logical agency; they may also be caused by aggregation of 
minuter vesicles, or by their bursting into each other ; or by 
protoplastic activities. Except where formed by direct proto- 
plastic ingestion, they are modifications of existing minuter 
inclusions; in all cases they are due to functions of the finer 
foam. Being products of a more primary set of structures 
and functions they are secondary in origin and may be purely 
incidental to the more intimate and basic life history and 
habit of the continuous substance. 

Once formed, each new series of vesicular inclusions neces- 
sarily introduces new interactions and new interrelations, 
chemical, physical and physiological. These can create for 
the mass such functional possibilities as may materially alter 
its habits as organism. For instance, while the finer foam 
carries what can for relatively short periods maintain its func- 
tional activities, a fluid structure of Butschli whose inclusions 
are of reserve similar material, can make the general or local 
substance independent of renewed appeal to external environ- 
ment for much longer periods. Such a structure by receiving 
excretions as well as excess of specific nutriment or stimulus 
can also prevent clogging, or impeding, or functional waste, of 
the active substance, and can save loss of time, force and sub- 
stance in deportation by interalveolar currents. It is a con- 
server of established habits as well as an initiator of a new 
series of functional relations. It offers increased leverage, and 
opportunity for organization of contractile powers on such a 
scale as can more patently alter shape or displace a mass or 
area. It keeps together relatively large quantities of sub- 
stances secreted or excreted by the general substance or pro- 

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1 64 ANDREWS. 

duced in situ, so that these, when poured out at intervals, can 
by their quantity strongly inlSuence other relatively large 
areas, or similar organs on an equal vesicular scale. It repre- 
sents naturally a certain decrease of osmotic compulsion, and 
of lamellar tension and mass viscosity, and greater economy of 
lamellar material with reservation of its stimulus. Its inclu- 
sions both by the physical opportunity and compulsions they 
offer, and by the chemical interactions they supply and create, 
serve indubitably to maintain the local and general substance 
in functional activity. Thus they come to mean reserves or 
grosser conditionings on which life and habit of the race sub- 
stance can depend. Such an alveolar structure alters and 
influences functional rhythms. Possibilities of substance para- 
sitism are much increased by it, and this is of radical impor- 
tance. Such a structure means again, increase of mass with 
many fruitful chemical and physical relations which can mater- 
ially affect the fate of organisms. Above all, it offers oppor- 
tunities for sudden irritable and contractile organization of 
specific sorts ; and for sudden intensification of the powers of 
the substance, as well as of swift recuperation and nutrition or 
stimulation for this, since formation of a finely subdivided foam 
from a structure of Butschli, brings at once into play an im- 
mense amount of reserve material and in such a manner as to 
)rield at a given moment stimulus and food to the largest area 
of irritable substance. Basis for organized response as well as 
nutrition and stimulus can thus be swiftly and enormously in- 
creased within a given space limit. Since this change takes 
place in the finer foam also, it is clear that even there, 
resources over and above those for immediate function are 
carried. I have already shown that formation and physical 
modification of ectosarcal areas must have a powerful effect on 
form, motions, and ingesting habit, besides exerting a radical 
guidance of the general habits and instincts, of organisms. 
Since a mere physical incident of local segregation of fluid- 
bearing alveoli, offers the contractile, irritable, substance op- 
portunity for organized function of various sorts, it seems the 
physical form alone of protoplasm may have been a potent, 
and even a cogent, factor in evolution of organisms along lines 

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leading into expansion, extension and intensification of the 
powers of the substance; — that is, it may have opened the 
way to, or even enforced, an ectosarcal set of substance habits. 
Much stress has been already laid upon formation and function 
of ectosarcal areas. They are indeed of almost unbounded 
importance in the history of the substance and organisms. 
Through these areas alone does the substance attain its most 
complete control of environment; through these alone it struc- 
turally reaches its higher life of the senses, of perception, of 
thought, of psychological growth and reach of function, all of 
which things are rooted in vesicular organization of irritability 
and contractility; — for in these areas the living substance is 
set free, fed, and stimulated, for fullest, highest and most 
characteristic self-expression. 

A single vesicle of the structure of Biitschli forms for the 
continuous substance a true organ, and when there are many 
whose interactions of causes and opportunities make up an 
organ, or set of functions, useful to the general substance or- 
ganism, — there has been formed within the mass a second- 
ary organism. However incidental their origin, each set of 
functional interrelations extends and conditions and makes 
more dependent the true primary substance organism, and, as 
we have known, may even obscure this as cause and end. In 
numerous ways, local substance organization could become 
responsive to internal or outside stimulus or influence. Over- 
nutrition, relative osmotic values of inclusions, dialysis as well 
as surface tension, — all these things are doubtless fruitful 
causes of origin and grouping of vesicular structures. Sub- 
stance habit is clearly guided and to some extent controlled by 
chemical and physical conditionings, and it is likely that of 
all such opportunities the living substance organism neglects 
in the long run no one that fits in well with its inherent or 
established lines of habit. It is to be kept well in mind that 
as far as we can go in the series, the continuous substance is 
still complex organism with specific controls and habits and 

The greater stability of the coarser structure c^ create and 
support more stable activities, form, functions, tendencies, and 

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1 66 ANDREWS. 

habits. In such a set of conditions the more patent organism 
has its origin. Now formation and function of substance 
organs being closely dependent on presence of specific mate- 
rials, their origin and meaning cannot be wholly inferred from 
immediately existing conditions, but must be taken in relation 
to race habits of the substance. Functions directly possible to 
such organs are complex enough, but that strange interwoven 
chemical, physical and physiological causation spoken of in 
Heredity multiplies others intermingled with these. What 
causative influence may come into each substance organ from 
any side and from long distances of space and time, for insti- 
gation or support of any local habit, especially in the invisible 
series, cannot be known or guessed. The excretions of one 
substance organ can stimulate secretive function in another, 
near or remote, this may be in turn necessary for excretions 
of another such organ, and these again cause physiological 
activity, as of contraction or increased irritability, in still 
others, the products of which again act or react functionally 
elsewhere. The products of one minute set of vesicular func- 
tions may stimulate organs of a structure of Butschli, and 
these again act upon grosser or compound organs, and each 
of all this series again react upon the primary substance or- 
ganization. Iterated or long continued function of one set 
may be necessary for more rarely recurrent, or even chance, 
functions of another substance organ. Chemical products of 
invisible as well as visible substance organs may be held 
sealed in vesicles of the finer foam, and later find their way, by 
migrant interalveolar stuff or by blood vessels, to a perpetuation 
or other area, there to await interactions of many other substance 
organs before their turn for causative function arrives. The 
whole organism may be regarded as a compound and complex 
series of many vesicular organs for excretion and secretion as 
well as for irritability or contractility. In this, and in the 
rhythms of recurrence of function, all sorts and kinds of Ata- 
vism are rooted. Every vesicle whose walls are of living func- 
tioning substance is a true substance organ. It is secretory 
and excretory for this which is muscle and nerve. A final 
hypothetical vesicle whose walls should be a simple film and 

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pel certain relations of the mass or organism with the opportuni- 
ties of external environment. It need no longer be regarded 
as meaning mental states, or as due to mental processes, for 
these, if present, are but part of the chain of interactions of 
substance organs. Apart from their incidental form as re- 
sults of secondary substance function and vesicular organiza- 
tion, — instincts are peculiarly grouped as those interrelations 
of substance habit that result in space-bridging and time-bridg- 
ing relations of the mass or animal with its external opportuni- 
ties. They involve always intensification areas of ectosarcal 
organization, — indeed, but for these, I think instincts as such 
would be wholly unknown to us, remaining indistinguishable 
amongst other substance habits within the mass, where doubt- 
less there are now hidden many kindred phenomena, having 
relation to the internal environment of the mass. For like all 
interrelations of the organism's parts, or of the mass with its 
external environment, instinct merely repeats in grosser terms 
radical powers and habits of the substance as such, expressed 
upon a secondary functional machinery. 


[151] To sum up; — the facts seem to warrant present belief 
that the living substance of all organisms is one physiologi- 
cally continuous, living, plasma, homogeneous throughout in 
its intrinsic powers and properties, but having varied local and 
temporary habits of self-expression, which are largely, and 
inextricably correlated with physical and chemical condition- 
ings of its form and composition as complex emulsive 
foam — yet not to be wholly identified with, nor wholly ex- 
plained by, these. The organism as we have known it, is 
secondary, incidental, to the life-history of the protoplastic con- 
tinuous substance of the living being: is result rather than 
cause of substance habit. Visible form and structure express 
only secondary truths. The part played in this state of things 
by purely physical and chemical conditions is doubtless enor- 
mous, but the wonder is not thereby lessened. If these re- 
sults, or the physical terms in which they are presented, have 

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at moments seemed to make the living organism ever more 
and more an automatic result of physical and chemical adjust- 
ments and relations; — let me remind, nay, urge upon the 
reader, that behind and beyond this automatism, the living 
substance has been shown to be still unrevealed to us, still 
transcending existing arrangements though always by and 
through these expressing its unexplained, inherent, powers and 
properties. For in all structures, however relatively stable 
they seem to us, the protoplastic substance is always ready 
and eager for more radical physiological self-expression. And 
it is this substance that is the true organism, by whose secre- 
tions, excretions, and dispositions of its surplus material in an 
organized vesicular way, the secondary and more patent organ- 
ism with which we are familiar is formed. Perhaps one of 
the most important things shown is that this protoplastic sub- 
stance is everywhere capable of just such organization of its 
powers in relation to its internal opportunities, as produces in 
organisms, from the lowest to the highest, all perceptive inter- 
course with mass environment. We may limit the word per- 
ceptive to the lowest form of sentience, but in and from this 
can arise by gradations the highest sorts known to us, even ap- 
perception and idea. In other words the substance organism 
has within the limits of invisible vesicular organization all that 
is requisite for true physiological habits and instincts, akin to 
those of the patent organism. The simplest and most primi- 
tive form of living substance we can get, and the smallest 
sub-divisions of this we can see with the microscope, is still a 
complex organism functioning for the race-substance, — is still, 
old as the hills, and very wise. 

In its inseparable physical form and chemical constitution 
lie the necessity and the temptation of a physical basis upon 
and through which to postulate the life phenomena of the 
living substance, since it expresses and must express itself 
through these conditionings. Because he has supplied in great 
part this intrinsic necessity of the science of biology, and in 
such form, that, with some extension and modification, it can be 
brought directly into relation with substance phenomena down 
to the smallest seen or inferred; — Biitschli deserves highest 

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gratitude and praise. Had it not been for his masterly work, 
these researches at least would not have been carried on so 
effectively, nor expressed in so unified a terminology which 
seems to add much to their value. I may say that, until after 
many repeated series of observations, I was quite neutral as to 
this vesicular theory, — if with any bias, was adverse to it. 
The possible interaction of physical, chemical, and physi- 
ological conditions in so sensitive, so powerful, and so causa- 
tive an arrangement as is found in the living emulsive 
foam, passes all imagination, and offers wonderful oppor- 
tunities of evolution along lines of substance habit. For 
another interesting result of these researches would seem 
to be a re-birth of "natural selection," making this again 
appear the powerful agent in evolution of organisms that Dar- 
win believed it to be. We must remember that he said, 
^* Natural Selection can act on every internal organ^ on every 
shade of constitutional differencCy on the whole machinery of 
life'' His line of argument becomes perforce artificial where- 
ever he attempts to decide the usefulness of any such differ- 
ence solely in relation to the organism in which it is 
found. Again and again, being confronted with this difficulty 
of linking values of structural or functional differences with 
the life-history of the organism, Darwin was compelled to find 
escape along lines of benefit to the species of organisms, — 
and this truly brings the matter at once into the ground of my 
offered standpoint. Natural selection acting upon substance 
structures and substance organs dives deeper into life mys- 
teries, is more searchingly constraining upon the race-history, 
than it ever could be by acting upon mere organism structures 
and organs, — that application of the theory must stumble and 
fail wherever it meets one of the many sacrifices of the unit for 
the substance organism as prime meaning or end. Instead of 
saying with Darwin " Natural selection it should never be for- 
gotten can act in each part of each being solely through and 
for its advantage," I would say rather, ' through and for the 
advantage of the substance as such and especially as race 
organ.' Re-reading the Origin of Species, I have been amazed 
at elimination of what have always seemed like obscurities 

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of reasoning and radical difficulties. Understanding of sexual 
selection and of the correlation of parts and powers in organ- 
isms has also been much helped for me by the new set of facts, 
as to selection of environment by the substance as such, and as 
to interrelation of substance organs in creating the organism, as 
well as in expressing the substance's protoplastic functional life. 

Reproduction is most helpful in race-history of the sub- 
stance not alone by multiplying the race organs, but by per- 
mitting the death of adult phases^ for thus it recurrently sets 
the substance free from trammels and limitations imposed 
upon it by that secondary machinery of the patent organism . 
its more intimate life and functions have created, which must 
limit its evolutionary progress as well as its continuance. 

The offered standpoint is a most fruitful one; yet it is not 
an easy thing at once to assume it, and for a time the mental 
foothold must feel slippery. Man is so used to regard as 
property of the animal, all structure and function within its 
limits. Belonging to a standpoint of early and almost inevit- 
able interpretation, and involving natural egotism of the unit, 
such mental custom is deep-rooted and difficult to overcome. 
A living being seems to be so patently master of its parts and 
powers, and these so clearly necessary to and for its existence; 
then too, — and here lies the root difficulty, — the coincidence 
of the external limits of the mass and of the animal, binds one 
to think of them as one and the same thing; the grosser 
structural subdivisions of the mass are plainly correlated with 
powers and functions of the animal we see. The simple, 
unconscious anthropomorphism of all our mental processes 
make this so natural a first standpoint, for how could the early 
self-conscious being escape a full and strong acceptance of 
himself as possessor and master of his parts and powers, — a 
belief that his directive influence in many ways, meant com- 
plete control. These things being true for himself, then the 
parts- and powers of every other animal must as surely express 
a like self-ownership and a kindred coherence of meaning for 
it as unit. 

Even the discovery of gross reflex actions, though a great 
shock to this primitive standpoint, has not displaced us from 

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it. And I think it will be long before most people will agree 
to think differently, before they will be willing to believe that 
all the structures and functions, visible and invisible, included 
in the skin of their familiar dog, or cat, or horse, do not pecu- 
liarly and wholly belong to that being, are not primarily for, 
and just to express, that individuality: — how otherwise ex- 
plain the strange unity and purposive harmony of external 
features, of grosser or even minuter structural characters, or 
the interrelations of parts and powers, and of functions; then 
again these things mean normal life and function to the organ- 
ism as such, deprivation of them means to it maimed or 
pathological states, or death even, and many of them are 
seen to act directly for and to subserve its existence. To ask 
man to think of these things as but incidental to minute, local, 
vesicular, organization and function of a protoplastic ground 
substance which must itself depend on that grosser organism 
for very continuance of life, — surely this is an unfair, even a 
ludicrous, demand upon human intelligence ! To even the 
thinking public, the question raised must for a long time to 
come seem to be mere mental juggling- — a sleight of idea 
trick, — or an effort to apply metaphysical methods and a mysti- 
cal standpoint to biological fact. If the sum total of parts 
and powers included within an animal's limits are not indeed 
its own, are not for it, — whose and for what are they ? Only 
three years back such a question would have seemed too absurd 
to be asked — may even yet seem so — and it is not many 
years since the means of perceiving or answering it have been 
in our hands. But I think to those, (they will be few), who 
will read with patient thought the long and minute record of 
selected facts in the foregoing pages, this question will not 
only thrust itself forward as it has done for me, but will bring 
with it an answer of sufficient assurance to swerve the trend 
of research more and more strongly toward the lines indicated. 
The offered standpoint does not miss the utmost interdepend- 
ence of parts and powers of any living unit, — it makes this 
more radically certain and clear than has yet been understood, 
showing that each and all are of value in the true organism's 
life; it does not hide from us that these form or maintain the 

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obvious creature, or that they are wholly dependent on it for 
continued existence, — but it discovers to us that this is but a 
small part of all, that the true organism is the invisible vesic- 
ular substance whose mass limits coincide of necessity with 
those of the living being; that all of the multiple parts and 
powers, functions and organs, of living units are, and primarily 
were, of and for the substance as such, and only partially and 
incidentally for the animal or plant : — in short that the zoolog- 
ical or botanical unit is in part a psychological formulation of 
mass separateness, while actually a vesicular accident or func- 
tional incident of the substance organism's protoplastic life. 

The fruit of these researches, then, is discovery of a new 
biological standpoint with its terms. Though bringing no 
explanation of vital phenomena, they may, none the less, prove 
helpful, — after the manner of a perpetuation area, — by freeing 
the substance of our thought from trammels of that secondary, 
incidental, structure its very self-expression has wrought about 
it ; and by re-embodying it in a younger and more plastic phase 
which, though it must bear out the race-history, may better 
meet our opportunities, and is less encumbered by excretions, 
and inorganic, inelastic, deposits of centuries of function. 

Our old standpoint will not, and should not, be abandoned 
for the new — but this, once perceived, can never be forgotten 
or discarded. Research cannot spring wholly from one or 
from the other, but the interest of active workers will surge 
into the new or back to old paths as natural limitations of 
power and focussed effort block man's progress here or there. 
Flux and intermission and recurrence — these things pertain 
to all substance function from the lowest protoplastic to the 
highest perceptive — marking the very throb and pulse of 

The mystery of life remains, — but at least it has had no 
light answer framed to meet it. " That which is is far oflf and 
exceeding deep ; who can find it out ? " 

August, 1896. 

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